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'ZS \A 




THIS BOOK I.IAY HOT BE PHOTOCOPIED 



PROCEEDINGS 



OF THE 



SCIENTIFIC MEETINGS 



OF THE 



ZOOLOGICAL SOCIETY 



OF LONDON 



FOE THE YEAE 



1887. 




FEINTED FOR THE SOCIETY, 
AND SOLD AT THEIR HOUSE IN HANOVER SQUARE. 

LONDON : 
MESSES. LONGMANS, GREEN, AND CO., 

PATERNOSTER ROW. 



LIST 



OF THE 



COUNCIL AND OFFICERS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON. 

1887. 



COUNCIL. 

{Elected April 29, 1887.) 

Peofessoe W. H. Flowee, C.B., LL.D., F.R.S., President. 



Liet7t.-Gen. The Loed AfirNGEE, 
C.B. 

William T. Blanfoed, Esq., 
P.R.S. 

Majoe-Gen. Heney Cleek, E.A., 
F.RS. 

Henet E. Deessee, Esq. 

Charles Deummond, Esq., Trea- 
surer. 

SiE Joseph Fateer, K.C.S.I., 
F.R.S., Vice-President. 

John P. Gassiot, Esq. 

Col. James A. Geant. C.B., C.iS.I., 
F.R.S. 

De. a. C. L. G. GiJNTHEE, 

F.R.S., Vice-President. 



Dr. Edward Hamilton, Vice- 
President. 

E. W. H. HoLDSwoRTH, Esq. 

Professor Mivarx, F.R.S., Vice- 
President. 

Professor Alfred Newton, 
M.A., F.R.S., Vice-President. 

Henht Pollock, Esq. 

The Lord Arthur Russell, 
Vice-President. 

Howard Saunders, Esq., F.L.S. 

Philip Lutley Sclatee, Esq., 
M.A.,Ph.D., F.R.S., Seeretcmj. 

Henry Seeboiim, Esq. 

Joseph Travers Smith, Esq. 

SuEG eon-Gen. L. C. Steavart. 



PRINCIPAL OFFICERS. 

P. L. Sclatee, Esc]., il.A., Ph.D., F.R.S., Secretar)/. 

Feank E. Beddard, Esq., M.A., Prosector. 

Mr. a. D. Bartlett, Superintendent of the Gardens. 

Me. F. H. Waterhouse, Librarian. 

Me. John Baeeow, Accountant. 

Mr. W. J. Williams. Chief Clerk. 



LIST 



OP THE 



OONTRIBUTOHS, 

With References to the several Articles contributed by each. 



Page 
Abraham, Rev. Nendick. 

On the Habits of the Tree Trapdoor Spider of Graham's 
Town 40 

Bartlett, a. D., Superintendent of the Society's Gardens. 

Remarks upon the Moulting of the Great Bird of Paradise. . 392 

Bates, H. W,, F.R.S. See Jacoby, Martin. 

Beddard, Frank E., M.A., F.R.S.E., F.Z S., Prosector to 
the Society, Lecturer on Biology at Guy's Hos[)ital. 

Notes on Brachyurus calvus. (Plate XII.) 119 

On the Structure of a new Genus of Lumbricidfe {Tham- 
nodrilus gulielmi) 1 54 

Contributions to the Anatomy of Earthworms. — Nos. I., 
II., III. (Plate XXXIII.) . . ." 372 

Note on, a Point in the Structure of Myrmecobius 527 

Contributions to the Anatomy of Earthworms. — No. IV. . ,544 

Notice of a Memoir entitled " Observations on the Structure 
of Hooker's Sea-Lion (Arctocephalus hooheri) " 640 



IV 



Page 



Bell, F. Jeffrey, M.A., Sec. R.M.S., F.Z.S., Professor of 
Comparative Anatomy in King's College, London. 

Exhibition of, and remarks upon, a bifid specimen of Nereis 
pelagica 3 

Report on a Collection of Echinodermata from the Anda- 
man Islands. (Plate XVI.) 139 

Extracts from a communication sent to him by Mr. Edgar 
Thurston, containing Observations on two Species of Batra- 
chians of the genus Cacopus 189 

Report on the Echinodermata collected by the Officers of 
H.M.S. ' Flying-Fish ' on Christmas Island 523 

Studies in the Holothuroidea. — VI. Descriptions of new 
Species. (Plate XLV.) 531 

Exhibition of, and remarks upon, the integumentary 
glands of a Rocky-Mountain Goat 579 

Bell, F. Jeffrey, M.A., Sec.R.M.S., F.Z.S., and Smith, 
Edgar A., F.Z.S. 

Observations on the " British Marine Area" 560 

BiDiE, Surgeon-Gen., C.I.E., C.M.Z.S. 

Exhibition of a photograph of, and remarks upon. Elephants 
in sexual congress 560 

Blanford, William Thomas, F.R.S., F.Z.S., &c. 

Critical Notes on the Nomenclature of Indian Mammals. . 620 

BOULENGER, G. A., F.Z.S. 

List of the Reptiles collected by Mr. H, H. Johnston on 
the Cameroons Mountain 127 

On a Collection of Reptiles and Batrachians made by Mr. 
H. Pryer in the Loo Choo Islands. (Plates XVII. & XVIII.) 146 

On a new Geckoid Lizard from British Guiana 153 

An Account of the Fishes collected by Mr. C. Buckley in 
Eastern Ecuador. (Plates XX.-XXIV.) 274 

Second Contribution to the Herpetology of the Solomon 
Islands. (Plate XXVIII.) 333 



V 

Page 
On a new Gecko, of the Genus Chondrodactylus, from 

the Kalahari Desert 339 

On a new Snake of the Genus Lamprophis now living in 
the Society's Gardens. (Plate XXXIV.) 397 

Eeport on the Reptiles collected by the Officers of H.M.S. 
' Flying-Fish ' on Christmas Island 516 

On the Systematic Position of the Genus Miolania, Owen 
(Cei-atochelys, Huxley) .554 

Notes on Emys blandingii. (Plate L.) 55.5 

Exhibition of, and remarks upon, some living specimens of 
an African Batrachian {Xenopus Icevis) 563 

A List of the Reptiles and Batrachians collected by Mr. 
H. H. Johnston on the Rio del Rey, Cameroons District, 
W. Africa 564 

On a new Species of Hyla from Port Hamilton, Corea, 
based on an example living in the Society's Gardens. 
(Plate LI.) 578 

Description of a new Genus of Lizards of the Family 
Teiidas 640 

An Account of the Fishes obtained by Surgeon-Major A. 
S. G. Jayakar at Muscat, East Coast of Arabia. (Plate LIV.) 653 

Brady, H. B., F.R.S., Parker, W. K., F.R.S., and Jones, 
T. Rupert, F.R.S. 

Notice of Memoir on the Foraminifera procured on the 
Abrohlos Bank during the Cruise of H.M.S. ' Plumper' . . 3 

Burmeister, Dr. H., F.M.Z.S. 

Letter from, describing a Humming-bird from Tucuman. . 638 

Butler, Arthur G., F.L.S., F.Z.S., &c. 

Report on the Lepidoptera collected by the Officers of 
H.M.S. ' Flying-Fish ' on Christmas Island 522 

On two small Collections of xlfrican Lepidoptera recently 
received from Mr. H. H. Johnston 567 



VI 

Page 

COCKERELL, T. D. A. 

Exhibition of, and notes upon, some Mollusca taken at 
Isleworth, Middlesex 362 

• Davis, James W., F.G.S. &c. 

Note on a Fossil Species of Clilamydoselachus 542 

Day, Francis, C.I.E., F.Z.S. 

Exhibition of a specimen of a hybrid Pilchard, and of a 
specimen of a Salmo j)urpuratus raised in this country .... 50 

On a supposed Hybrid between the Pilchard {Chtpea pil- 
chardus) and the Herring (C. harengus)^ and on a specimen 
of Salmo purpuratus. (Plate XV.) 129 

On the Occurrence of Scorpcena scrofa off the South Coast 
of England 342 

Dendy, Arthur, B.Sc, F.L.S. 

Abstract of a Memoir on the West-Indian Chulinma;, with 
Descriptions of new Species 503 

Report on the Porifera collected by the Officers of H.M.S. 
' Flying-Fish ' on Christmas Island. (Plate XLIV.) 524 

DoBsoN, G. E., M.A., F.R.S., F.Z.S., &c. 

On the Genus Myosorex, with Description of a new Species 
from the Rio del Rey (Cameroons) District 575 

Douglas-Ogilby, J. See Ogilby, J. Douglas. 

Dresser, Henry E., F.Z.S. 

Exhibition of, and remarks upon, some specimens of a Tit- 
mouse obtained by Dr. Guillemard in Cyprus 5G3 

Druce, Herbert, F.L.S., F.R.G.S., F.Z.S. 

Descriptions of some new Species of Lepidoptera Hetero- 
cera, mostly from Tropical Africa. (Plate LV.) 608 

Elwes, Henry John, F.Z.S. 

Description of some new Lepidoptera from Sikkim 444 



Page 

EVBRETT, A., C.M.Z.S. 

Extract from a letter, containing a notice of the Collection 
made by Mr. "Whitehead on Kina Balu Mountain in Nor- 
thern Borneo 502 

FisK, Rev. George H. R., C.M.Z.S. 

Extract from a letter, respecting the killing and eating, 
by a Mouse, of a young venomous Snake 340 

Flower, William Henry, C.B., LL.D., F.R.S., F.L.S., 

President of the Society, Director of the Natural History 
Collection, British Museum. 

Extracts from a letter from Dr. Emin Pasha, containing 
some remarks upon Natural History specimens procured in 
Moubottu 468 

Exhibition of a photograph of, and remarks upon, a 
specimen of Rudolphi's Whale {Balcenoptera borealis) taken 
in the Thames 564 

On the Pygmy Hippopotamus of Liberia, Hippopotamus 
liberiensis (Morton), and its claims to distinct Generic Rank 612 

Garrett, Andrew, of Huahine, Society Islands. 

On the Terrestrial Mollusks of the Viti Islands.— Part I. 164 
On the Terrestrial Mollusks of the Viti Islands.— Part II. 284 

Giglioli, Henry H., C.M.Z.S., and Salvadori, Thomas, 
C.M.Z.S. 

Brief Notes on the Fauna of Corea and the adjoining coast 
of Manchuria. (Plate LII.) 580 

GoDMAN, Frederick DuCane, F.R.S., F.L.S., F.Z.S. See 
Sal\in, Osbert, F.R.S. 

Gorham, Rev. H. S., F.Z.S., F.E.S. 

On the Classification of the Coleoptera of the Subfamily 
Languriides 358 

Revision of the Japanese Species of the Coleopterous Family 
EndomychidtB. (Plate LIII.) 642 



vin 

Page 

Gkant, W. R. Ogilvik. 

A List of tlie Birds collected b}' Mr. Charles Morris 
Woodford in the Solomon Archipelago. (Plate XXVIl.) . . 328 

GiJNTHER, Albert, C. L. G., M.A., M.U., Ph.D., F.R.S., 
V.P.Z.S., Keeper of the Zoological Department, British 

Museum. 

Exhibition of a hybrid between a Golden Plieasant and a 
Reeves' Pheasant, also of a hybrid between a white Fantail 
Pigeon and a Collared Dove 503 

Report on a Zoological Collection made by the Officers of 
H.M.S. 'Flying-Fish' at Christmas Island, Indian Ocean. 
(Plates XLI.-XLIV.) 507 

Descriptions of two new Species of Fishes from Mauritius. 
(Plates XLVIII. & XLIX.) 550 

Howes, G. B., F.Z.S., F.L.S,, Assistant Professor of Zoology, 
Normal School of Science and R. School of Mines, S. 
Kensington. 

On the Skeleton and Affinities of the Paired Fins of Cera- 
todus, with Observations upon those of the ElasmobrancJdi. 
(Plates I.-III.) 3 

Exhibition of, and remarks upon, an original drawing of 
the head of an abnormal Palinurus (P. penicillatus) 4G8 

On a hitherto unrecognized Feature in the Larynx of the 
Anurous Amphibia 491 

Hume, A. O., C.B., F.Z.S. 

Remarks on certain Asiatic Ruminants. — I. Budorcas 
taxkolor, Hodgson. The Gnu-goat or Takin 483 

Jacoby, Martin, F.E.S. 

Descriptions of the Phytophagous Coleoptera of Ceylon, 
obtained by Mr. George Lewis during the years 1881-82. 
(Plates X. & XL) C5 



IX 

Page 
List of a small Collection of Coleoptera obtained by Mr. W. 

L. Sclater in British Guiana. With the Description of a 

new Species by H. W. Bates, F.R.S 490 

Jenner-Weir, J. See Weir, John Jenner. 

Jones, T. Rupert, F.R.S. &c. See Brady, H. B. 

KiRBY, W. F., F.E.S., Assistant in the Zoological Department, 
British Museum. 
Notice of a Memoir on the Subfamily Libellulince 470 

Leech, John Henry, B.A., F.R.G.S., F.L.S., F.Z.S., &c. 

Exhibition of specimens of some new Butterflies from Japan 
and Corea , 342 

On the Lepidoptera of Japan and Corea.— Part L Rho- 
palocera. (Plates XXXV, & XXXVL) 398 

Le Souef, Albert A. C, C.M.Z.S. 

Letter from, containing remarks upon some living Duck- 
bills (Or/it7Aor%nc/«M5jt3a?-ac?o^Ms) in confinement at Melbourne 363 
Maclear, Captain. 

Report on Christmas Island 5q8 

Menzbier, Dr. M., C.M.Z.S., Professor in the University of 
Moscow. 

On a new Caucasian Goat {Capra severtzowi, sp. n.) 618 

Nathusius, W. von. 

Exhibition of, and remarks upon, some preparations of 
Symbiotes k - n 

Newton, Alfred, M.A., F.R.S., F.L.S., F.Z.S., Professor of 
Zoology and Comparative Anatomy in the University of 
Cambridge. 

Exhibition of, and remarks upon, a specimen of Bulwer's 
Petrel found in Yorkshire ggo 

NicfviLLE, Lionel de, F.E.S. 

Descriptions of some new or little-known Butterflies from 
India, with some Notes on the Seasonal Dimorphism obtaining 
in the Genus Melanitis. (Plates XXXIX. & XL.) T 448 



X 

Page 
Ogilby, J. Douglas, Ichthyological Department, Australian 

Museum, Sydney. 

Description of a little-known Australian Fish of the Genus 
Oirella 393 

On an undescribed Fish of the Genus Prionurus from 
Australia 395 

On a new Genus and Species of Australian Mugilidae. . . . 614 

On a new Genus of Percidae 616 

Ogilvie-Grant, W. R, See Grant, W. R. Ogilvie. 

Parker, T. Jeffery, B.Sc, C.M.Z.S., Professor of Biology 
in the University of Otago, New Zealand. 
Notes on Carcharodon rondeletii. (Plates IV.-VIII.) . . 2/ 

Parker, T. Jeffery, B.Sc., C.M.Z.S., and Scott, Professor 
J. H. 
Notice of a memoir on a Whale of the Genus ZipMus, 
recently obtained near Dunedin, New Zealand 342 

Parker, W. K., F.R.S., &c. See Brady, H. B. 

Pasha, Dr. Emin, C.M.Z.S. 

Letter from, referring to some Collections sent to the 
British Museum 564 

Phipson, H. M., C.M.Z.S. 

Letter from, concerning living specimens of two Snakes . , 639 

PococK, R. Innes. 

Report on the Crustacea collected by the Officers of 
H.M.S. ' Flying-Fish ' on Christmas Island 520 

PouLTON, E. B., M.A., F.Z.S., F.L.S., of Jesus and Keble 

Colleges, Oxford, Lecturer on Zoology and Comparative 

Anatomy, St. Mary's Hospital, Paddington. 

The Experimental Proof of the Protective Value of Colour 

and Markings in Insects in reference to their Vertebrate 

Enemies 191 



Page 
Salvadori, Thomas, C.M.Z.S., anil Giglioli, Henry H., 
C.M.Z.S. 

Brief Notes on the Fauna of Corea and the adjoining 
coast of Manchuria. (Plate LII.) 580 

Salvin, Osbert, F.R.S., F.Z.S. 

Exhibition, on behalf of Mr. F. D. Godman, and remarks 
upon, a pair of Ornithoptera victories 190 

Saunders, Howard, F.L.S., F.Z.S., &c. 

Exhibition of, and remarks upon, a specimen of the Medi- 
terranean Black-headed Gull {Larus melanocephalus), shot 
near Great Yarmouth 2 

Exhibition of, and remarks upon, a Harlequin Duck {Cos- 
monetta histrionica) shot near the Fame Islands 319 

Exhibition of, and remarks upon, a specimen of Saccicola 
isabellina, shot in Cumberland 579 

Sclater, Philip Lutley, M.A., Ph.D., F.R.S., Secretary to 
the Society. 

Report on the additions to the Society's Menagerie in 
December 1886 1 

Exhibition of, and remarks upon, a specimen of a rar 
Parrot, Chrysotis bodini 2 

Characters of new Species of Birds of the Family Tyran- 
nidse. (Plate IX.) 47 

Report on the additions to the Society's Menagerie in 
January 1887 138 

Report on the additions to the Society's Menagerie in 
February 1 887 319 

Report on the additions to the Society's Menagerie in 
March 1887 - 340 

Extracts from a letter addressed to him by the Rev. Geo. 
H. R. Fisk, C.M.Z.S., respecting the killing and eating, by 
a Mouse, of a young venomous Snake 340 



xii 

Page 

Exhibition of, and remarks upon, a set of eleven photo- 
graphs, representing objects of Natural History collected by 
General Prejevalski in Central Asia 362 

Extract from a letter addressed to him by Mr. Albert A. C. 
Le Souef, C.M.Z.S., containing remarks upon some living 
Duckbills iOrnithorhynchus paradoxus) in confinement at 
Melbourne 363 

Report on the additions to the Society's Menagerie in 
April 1887 396 

Extracts from a letter addressed to him by Mr. Roland 
Trimen, F.Z.S., respecting the obtaining of a second example 
of Laniarius atrocroceus 396 

Report on the additions to the Society's Menagerie in 
May 1887 482 

Remarks upon specimens of two Species of North- American 
Foxes living in the Society's Gardens 482 

Exhibition of, and remarks upon, the skin of a White- 
nosed Monkey (Cercopithecus ascanias ?) , 502 

Exhibition of, and remarks upon, a specimen of a Pheasant 
from Northern Afghanistan (^Phasianus principalis) 502 

Note on the Wild Goats of the Caucasus 552 

Report on the additions to the Society's Menagerie in June, 
July, August, September, and October, 1887 558 

Report on the additions to the Society's Menagerie in 
November 1887 638 

Extract of a letter from Dr. Burmeister, F.M.Z.S., and 
remarks on a supposed new Humming-bird (Chceioeercus 
burmeisteri) 638 

ScLATER, W. L., B.A., F.Z.S. 

Notes on the Peripatus of British Guiana 130 

Scott, Prof. J. H., and Parker, Prof. T. Jeffery. 

Notice of a memoir on a Whale of the Genus Ziphius, 
recently obtained near Dunedin, New Zealand 342 



xm 

Page 
Seebohm, Henry, F.L.S., F.Z.S., &c. 

Description of a supposed new Species of the Genus Merula 
from South America ^^7 

Sharpe, R. Bowdler, F.L.S., F.Z.S., &c., Department of 
Zoology, British Museum. 

On a second Collection of Birds formed by Mr. L. Wray in 
the Mountains of Perak, Malay Peninsula. (Plates XXXVII. 
&XXXVIII.) 431 

Notes on Specimens in the Hume Collection of Birds. — 
No. 5. On Syrnium maingayi 470 

Report on the Birds collected by the Officers of H.M.S. 
' Flying-Fish ' on Christmas Island. (Plate XLIII.) 515 

On a new Species of Calyptomena 558 

Shelley, Capt. G. E., F.Z.S. 

On a Collection of Birds made by Mr. H. H. Johnston in 
the Caraeroons Mountain. (Plates XIII. & XIV.) 122 

SHUFELDT, R. W., M.D., C.M.Z.S. 

Notes on the Visceral Anatomy of certain Auks 43 

Smith, Edgar A., F.Z.S. 

On the Mollusca collected on the Cameroons Mountain by 
Mr. H. H. Johnston 127 

Notes on a small Collection of Shells from the Loo Choo 
Islands 316 

Report oii the MoUusks collected by the Officers of 
H.M.S. ' Flying-Fish ' on Christmas Island 517 

Notes on three Species of Shells from the Rio del Rey, 
Cameroons 566 

Smith, Edgar A., F.Z.S., and Bell, Prof. F. Jeffrey, 
F.Z.S. 

Observations on the "British Marine Area " 560 



XIV 

Page 
Styan, F. W., F.Z.S. 

Exhibition of a series of eggs of Chinese birds 2 

Sutton, J. Bland, F.R.C.S., Lecturer on Comparative 
Anatomy, Middlesex Hospital, Erasmus Wilson Lecturer 
on Pathology, Royal College of Surgeons. 

On some Specimens of Disease from Mammals in the 
Society's Gardens 364 

On the Arm-glands of the Lemurs 369 

Symonds, Edmond, of Kroonstad, Orange Free State. 

Notes on some Species of South-African Snakes 486 

Taczanowski, L., C.M.Z.S. 

Liste des Oiseaux recueillis en Coree par M. Jean 
Kahnowski 596 

Tegetmeier, W. B., F.Z.S. 

Exhibition of, and remarks upon, some heads of the 
Sumatran Rhinoceros, from Sarawak, Borneo 3 

Thomas, Oldfield, F.Z.S. 

List of Mammals from the Cameroons Mountain, collected 

by Mr. H. H. Johnston 121 

On the small Mammalia collected in Demerara by Mr. W. 
L. Sclater. (Plate XIX.). . 150 

On the Bats collected by Mr. C. M. "Woodford in the 
Solomon Islands. (Plates XXV. & XXVI.) 320 

On the Milk-dentition of the Koala 338 

Report on the Mammalia collected by the X)fficers of 
H.M.S. ' Flying-Fish ' on Christmas Island. (Plates XLI. 
&XLII.) 511 

Thomson, Arthur. 

Report on the Insect-house for 1886 50 

Exhibition of, and remarks upon, specimens of Papilio 
porthaon reared in the Society's Insect-house 4C8 



XV 

Page 
Trimen, Roland, F.R.S., F.Z.S., &c. 

Letter from, respecting the obtaining of a second example 
of Laniarius atrocroceus 396 

On Bipalium kewense at the Cape 548 

"Waterhouse, Charles O., F.E.S. 

On some Coleopterous Insects collected by Mr. H. H. 

Johnston on the Cameroons Mountain 128 

» 
Note on a new Parasitic Dipterous Insect of the Family 

Hippoboscidse 163 

Report on the Coleoptera collected by the Officers of 
H.M.S. ' Flying-Fish ' on Christmas Island 520 

Weir, John Jenner, F.Z.S. 

Exhibition of, and remarks upon, a skull of a feral Boar 
{Sus scrofa), obtained at Tauranga, New Zealand 397 

WiNDLE, Bertram C. A., M.A., M.D. (Dubl.), Professor of 
Anatomy in the Queen's College, Birmingham. 
On the Anatomy of Hydromys chrysogaster 53 

"Woodward, A. Smith, F.Z.S., F.G.S., of the British Museum 
(Natural History). 
On the Presence of a Canal-system, evidently Sensory, in 
the Shields of Pteraspidian Fishes 478 

Note on the " Lateral Line " of Squaloraja 481 

On the Fossil Teleostean Genus Rhacolepis, Agass. (Plates 
XLVI. & XLVII.) 535 

Wray, Richard S., B.Sc. (Lond.). 

Note on a Vestigial Structure in the i\.dult Ostrich 
representing the Distal Phalanges of Digit iii 283 

On some Points in the Morphology of the Wings of Birds. 
. (Plates XXIX.-XXXII.) 343 

Yerbury, Major, C.M.Z.S. 

Exhibition of, and remarks upon, a pair of horns of the 
Oorial 639 



LIST OF PLATES. 

1887. 



Plate 

I. 

II. 

III. 

IV. 

V. 

VI. 

VII. 

VIII. 

IX. 

X 

XI 

Xll. 

XIII. 

XIV. 

XV. 

XVI. 

XVII. 

XVIII. 

XIX. 

XX. 

XXI. 



Fins of Ceratodus and Protopferus 

Pelvic fins of Ceratodus 

Fins of Elasmobranchs and PolyiJterus 

Figs. 1 and 3. Carcharodon rondeletii. Figs. 2 and 4. 

Lamna cornuhica 

Figs. 5-10. Carcharodon rondeletii. Fig. 11. Lamna 

cornubica 

Carcharodon rondeletii 

Carcharodon rondeletii 

Carcharodon rondeletii (foetus) 



Page 

3 



27 



Fig. 1. Euscarthmus apicalis. Fig. 2. Leptopogon 



oustaleti 



\ Phytophagous Coleoptera of Ceylon 



47 
119 



XXII. 
XXIII. 

XXIV. 

XXV. 

XXVI. 



Brachyurus calvus 

Laniarius atroflavus ] 

Fig. 1. Zoster ops melanocephala. Fig. 2. Ploceus\ 122 

melanogaster 

Hybrid between Pilchard and Herring 

Echinoderms of the Andamau Islands 

Fig. 1. Japalura polygonata. Fig. 2. Taehydromus 

smaragdinus 

Fig. 1 . Taehydromus smaragdinus. Fig. 2. Lygosoma 

pellopleurum. Fig 3. Tropidonotas pryeri 

Hesperomys (Rhipidomys) sclateri 

Fig. 1. Pimelodus buckleyi. Fig. 2. P. longicauda. . 
Fig. 1. Pimelodus pule her. Fig. 2. Stygogenes hum- 

holdtii. Fig. 3. Nannoglanis fasciatus. Fig. 4. 

Stegophilus punctatus 

Chcetostomtis dermorhynchus '. 

Fig. 1. Parodon buckleyi. Fig. 2. Piabucina elon- 

gata. Fig. 3. Leptagoniutes steindachneri 

Sternarchus curvirostris J 

Pteropus grandis ( 

Nesoiiycteris woodfordi ■ ( 

6 



12-) 
139 

146 

150 

274 



320 



Plate 

XXVII. 
XXVIII. 



XXIX. 
XXX. 

XXXI. I 
XXXII. 
XXXIII. 
XXXI V. 

XXXV. 

XXXVI. 

XXXVII. 

XXXVIII. 

XXXIX. 

XL. 

XLI. 

XLII. 

XLTIJ. 

XLIV. 

XLV 

XLVI 

XLVII 

XLVIII 

XLIX, 

L 

LI 

LII 

LIII 

LIV 



:( 



LV. 



XVIU 

Page 

Macrocorax woodfordi 328 

Fig. 1. Lepidodactylus woodfordi. Fig. 2. Typhlops 
aluensis. Fig. 3. Batrachylodes vertebralis. Fig. 4. 
Hyla lutea 333 

Morphology of the Wings of Birds 343 

Anatomy of Eudrilus sylvicola 372 

Lahiprophis jiski 397 

Fig. 1. Papilio mikado. Fig. 2. Apatura cauta. Fig. 3. j 

cJ 2 Polyommatus auratus [■ 398 

Fig. L (S 2 Eiiripus coreanus. Fig 2. Dipsasflamen.\ 

Trochcdopterum peninsula ( joi 

Fig. 1. Minla soror. Fig. 2. jEthopyga wrayi .... 1 

New Indian Butterflies 448 

Pteropus natalis \ 

Mus macleari / 

Carpophaga whartoni T 

Packychalina spinosissima ) 

Characteristic spicules of new Holothurians 531 

Rhacolepis 535 

Latilus fronticinctus ■ I ckq 

Platycephalus suhfasciatus I 

Fig. 1. Emys blandingii. Fig. 2. Emys orbicularis. . . . 555 

Fig. 1. Hyla stepheni. Fig. 2. Hyla arborea,vsir.japonica. 578 

Fig. 1. Cygnus davidi. Fig. 2. Cyynus bewicki 580 

New Japanese Species of Endomychida 642 

Fig. 1. Opisthognathus muscatensis. Fig. 2. Gobius 

jayakari 653 

New African Lepidoptera 668 



LIST OF WOODCUTS. 

188/. 



Page 

Anterior aspect of the lower larynx, heart, and viscera of Synthli- 

borhamplms antiquus 45 

Anterior aspect of the lower larynx, heart, and viscera of Braclnj- 

rhamphus marmoratus ^ 45 

Cervical and brachial nerves of Hydromys 59 

Lumbar and sacral nerves of Hydromys 60 

Left hand and foot of Hydromys chrysoyaster . . 61 

Upper and lower incisors of Hydromys chrysoyaster 61 

Upper and lower jaw of Hydromys chrysoyaster 62 

Upper molar of Hydromys chrysoyaster 62 

Stomach of Hydromys chrysoyaster , 62 

CiBcum of Hydromys chrysoyaster 63 

Superior surface of liver of Hydromys chrysoyaster 64 

Under surface of liver of Hydromys chrysoyaster 64 

Gibhus {EdentuUna) johnstoni 128 

Gonatodes annularis 153 

Thamnodrilus yulielmi, from the ventral surface 155 

Thatnnodrilus gidielmi, anterior segments seen laterally 157 

Thamnodrilus yulielmi, seta from one of the segments of the clitellum 157 

Chief Trunks of Vascular System of Thamnodrilus 158 

One of the Anterior Nephridia of Thamnodrilus 160 

One of the Posterior Nephridia of Thamnodrilus 161 

Anapera Jhnhriata 163 

Phalanx 1 and the vestigial cartilage of digit iii., adult Ostrich .... 284 
The distal part of digit in. in the manus of the embryo of Ostrich . . 284 

Ventral view of left manus of embryo of Ostrich 284 

Helix largillierti, var 317 

Teeth of Pteropus grandis 321 

Skull of Nesonycteris woodfordi, upper view 325 

Skull of Nesonycteris ivoodfordi, side view 325 

Head of young Koala, showing milk-dentition 338 

Preparations of the distal cubital remiges, with their attached 

tectrices majores, of the Pheasant 346 

Preparations of the distal cubital remiges, with their attached 

tectrices majores, of the Golden Eagle 346 

Preparations of the manus of the Ostiich, showing the primaries and 

the manner of their attachment to the bones 351 



XX 

Page 
Dorsal view of the autebrachiuoi and maims of the wing of the 

Ostrich 351 

Ventral view of the manus of an embryo Ostrich 351 

Transverse section of the thorax of a rickety Monkey 364 

Under view of the skull-vault of a rickety Lion 365 

A longitudinal section of the head of a Lion-cub, showing over- 
growth of the tentorium cerebelli, &c 366 

Abnormal growth of papilla on the hind feet of a Coati 368 

Two Goat's feet with overgrown hoofs 368 

The forearm of Hapakmur griseus, showing the patch of spine-like 

processes and the tuft of hairs 369 

The forearm of Chirogaleus coquereli, showing the tuft of long hairs . 370 
The forearm of Lemur caftn, showing the raised patch of hairless 

skin covering the collection of sweat-ducts 370 

ForeaiTu of an adult Lemur catta, showing the blunt spur 371 

Arm of a fostal Lemur catta, to show the tuft of long hairs ........ 371 

Dissection of genital region of Eudrihis sylvicola 381 

Acanthodrihis dissimilis. Dissection of genital region 388 

Clitellar seta of Perichata houlkti 390 

Ce'^hdlonoi Palinurus penicillatus, \M&\-\mf an antenniform ophthalmite 469 

Fragmentary Median Plate of Shield of Pferaspis crouchii 479 

Horns of Budorcas taxicolor 484 

Phileurus sclateri 490 

The larynx in Rana esculenta and P. femporaria 492 

The larynx in Leptodactyhis pentadactylus 494 

The larynx in Rana 2)ipiens 494 

The larynx in Ceratophrys americana and C'alyptocephalus gayi .... 495 

The larynx and floor of the mouth in Chiroleptes australis 497 

The larynx in Chiroleptes anstralis, side view 498 

The larynx in Chiroleptes australis, longitudinal section 498 

Succinea solidula and <S'. solitaria 518 

Littorina granicostata 619 

Piezonoius discoidalis 521 

Vadebra inacleari 522 

Terias amplcia 523 

Pachychaliiia spinosksima 525 

Under surface of head of Mynnecohius fasciatus 627 

Glandular patch of My rmecohius fasciatus 528 

Minute structure of glandular patch of Myrmecobius 530 

Ciyptodrilus fletcheri 547 

Skull of Myosorex johnstoni 577 

Chthamalopteryx melbouniensis 616 

Stenolepis ridleyi 641 



PROCEEDINGS 



OF THE 



SCIENTIFIC MEETINGS 



OF THE 



ZOOLOGICAL SOCIETY 



OF LONDON 



FOE THE YEAR 



1887. 



(PLATES.) 




PRINTED FOR THE SOCIETY, 
AKD SOLD AT THEIR HOUSE IN HANOVER SQUARE. 

LONDON : 
MESSRS. LONGMANS, GREEN, AND CO., 

PATERNOSTER ROW. 



LIST OF PLATES. 

1887. 



Plaltf 

I. 

II. 

III. 

IV. 

V. 

VI. 

VII. 

VIII. 

IX. 

X.I 
XI. I 
XII. 
XIII. 
XIV. 

XV. 

XVI. 

XVII. 

XVIII. 

XIX. 

XX. 

XXI. 



XXII. 
XXIII. 

XXIV. 

XXV. 

XXVI. 



Fins of Ceratoihis and Prot.opterus 

Pelvic fins of C'eratodus 

Fins of Elasmobranchs and Polypterus 

Figs. 1 and 'A. Carchnrndnn roiiilelelii. Figs. 2 and 4. 

Lnmna cornuhica 

Figs. 5-10. Corctiurodonrondeletii. Fig. 11. Lamnn 

cornvbica f 

Carclwodon rondeletii I 

Carcharodon rondeletii I 

Carcharodon rondeletii (foetus ) ' 

Fig. 1. Euscarthmus apiccdis. Fig. '2. Leptopoyon 

oustaleti 

Phytophagous Coleoptera of Ceylon 






ens '• 



BracJiyurus calvus 

Laniariiis atroflariis 

Fig. 1. Zosterops melanocephala. Fig. 2. Ploce 

melonogaster ' 

Hybrid between Pilchnrd and Herring 

Echinoderms of the Andaman Islands 

Fig. 1. Japaliira polygonata. Fig. 2. Tachydromus \ 

smaragdinus / 

Fig. 1 . Tachydromus smaragdinus. Fig. 2. Lygosoma ( 

pellopleurum. Fig. 3. Tropidonofus pryeri ) 

Hesperomys [Rkipidomys] sclateri 

Fig. I. Pimelodus buckleyi. Fig. 2. P. longicauda. . ~ 
Fig. 1. Pimelodus pulcher. Fig. 2. Stygogenes hum- 

holdtii. Fig. 3. Nannoglanis fasciatus . Fig. 4. 

Stegophihis punctatus 

Cheetostomus dermorhynchus ' 

Fig. 1. Pnrodon buckleyi. Fig. 2. Pinbiicina elon- 

gata. Fig. 3. Leptagoniates steindachiieri 

Sternarchus curvirostris J 

Pteropus grandis I 

Nesonycteris woodfordi ( 



47 
«.") 

11!» 

122 

]2<> 
139 

146 

150 



320 



Plate 

XXVII. 
XXVIII. 




XXXVI. 

XXX VI I. 

XXXVIII. 

XXXIX. 

XL. 

XLI. 

XLII. 

XLTII. 

XLIV. 

XLV. 

XLVI. 

XLVII. 

XLVIII. 

XLIX. 

L. 

LI. 

LII. 

LIII. 

LIV. 

LV. 



Page 

Macrocorax woodfordi 328 

Fig. 1. Lepidodactylus woodfordi. Fig. 2. Typhlops 

aluetisis. Fig. 3. Batrachylodes vertehralis. Fig. 4. 

Hyia lutea 333 

Morphology of the Wings of Birds 343 

Anatomy of Eudrilus sylvicola 372 

Lamprophis fiski 39^ 

Fig. I. Papilio mikado. Fig. 2. Apatura cauta. Fig. 3. ■ 

(5' $ Polyommatus auratus \ 398 

Fig. 1. (S 2 Euripus coreanus. Fig- 2. Dipsasflamen.) 

Trochalopterum peninsulce I ^^ji 

Fig. 1. Minlu soror. Fig. 2. jEthopyga wrayi . . . . i 

New Indian Butterflies 448 

Pteropus natnlis \ 

Mus inacleari / _ 

Corpophaya whartoni 1 

I'achychalina spinosissima . ) 

Characteristic spicules of new Holothurians , . . . 531 

Rhacolepis 535 

Latilus fronlicinctiis I ^^q 

Platyceplialus subfasciatus I 

Fig. 1. Emys hlandingii. Fig. 2. Emys orbicularis. . . . 655 
Fig. L Hylastepheni. Fig. 2. Hyla arborea,va.r.japonica. 578 

Fig. 1. Cygnus davidi. Fig. 2. Cyynus bewicki 580 

New Japanese Species of Endomychida 642 

Fig. 1. Opisthognathus muscutemis. Fig. 2. Gobius 

jayaJcari "5o 

New African Lepidoptcra 668 



l.IST OF THE PUBLICATIONS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON. 



The scientific publications of the Zoological Society are of two 
kinds — " Proceedings," published in an octavo form, and 
" Transactions," in quarto. 

According to the present arrangements, the "Proceedings" 
contain not only notices of all business transacted at the scien- 
tific meetings, but also all the papers read at such meetings 
and recommended to be published in the " Proceedings " by the 
Committee of Publication. From fifty to seventy coloured plates 
and engravings are attached to each annual volume of the 
" Proceedings," to illustrate the new or otherwise remarkable 
species of animals described in them. Amongst such illustra- 
tions, figures of the new or rare species acquired in a living 
state for the Society's Gardens are often given. 

The "Proceedings" for each year are issued in four parts, on the 
first of the months of June, August, October, and April, the part 
published in April completing the volume for the preceding 
year. The price is 12*. per part for the edition vi^ith coloured, 
and 3s. per part for that with uncoloured Plates, 

The "Transactions" contain such of the more important 
communications made to the scientific meetings of the Society 
as, on account of the nature of the plates required to illustrate 
them, are better adapted for publication in the quarto form. 
They are issued at irregular intervals. 

Fellows and Corresponding Members, upon payment of 
a Subscription of One Guinea before the day of the Anni- 
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Society's Publications for the year. They are likewise entitled 
to purchase the Publications of the Society at 35 per cent, less 
than the price charged for them to the Public. A further re- 
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issued prior to 1861, if they exceed the value of five pounds. 

The following is a complete list of the publications of the 
Society already issued. They may be obtained at the Society's 
Office (3 Hanover Square, W.), at Messrs. Longmans', the 
Society's publishers (Paternoster How, E.G.), or through any 
bookseller. 
[June 1, 1887.] 



TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON. 

4to. 11 vols, and Six Parts. 



Vol. I., containing 59 Plates 
Vol. II., „ 71 „ 

Vol. III., „ 63 „ 
Vol. IV., „ 78 „ 

Vol. v., „ 67 „ 
Vol. VI., „ 91 „ 
Vol. VII., „ 73 „ 

Vol. VIII., „ 82 „ 
Vol. IX., „ 99 „ 

Vol. X., „ 94 „ 

Index, Vols. I.-X 

Vol. XI., part 1, containing 4 Plate 



(1833-35) 
(1835-41) 
(1842-49) 
(1851-62) 
(1862-66) 
(1866-69) 
(1869-72) 
(1872-74) 
(1875-77) 
(1877-79) 



To FellowB. 
j^ s. d. 
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Vol. XL, 
Vol. XL, „ 
VoLXL, „ 
VoLXL, „ 
Vol. XL, „ 
Vol. XL, „ 
Vol. XL, „ 
VoLXL, „ 
VoLXL, „ 
Vol. XL, „ 
Vol. XII., „ 
Vol. XII., „ 
Vol. XII., „ 
Vol. XIL, „ 
Vol. XIL, „ 
Vol. XIL, „ 



2, 
3, 
4, 
5, 
6, 
7, 
8, 
9, 
10, 
11, 
1, 
2, 
3, 
4, 
6, 
6, 



3 

13 

6 

9 

11 

10 

12 

14 

6 

7 

2 

12 

5 

7 



s (Jan. 
(Aug. 
(Mar. 
(Apr. 
(June 
(Jan. 
(Oct. 
(Jan. 
(Oct. 
(May 
(Nov. 
(Feb. 
(Apr. 
(Aug. 
(Oct. 
(Dec. 
(Apr. 



1880) 

1880) 

1881) 

1881) 

1881) 

1882) 

1882) 

1883) 

1883) 

1885) 

1885) 

1886) 

1886) 

1886) 

1886) 

I'^se) 

1887) 



2 
4 
5 

8 17 

9 8 
12 
10 3 

7 6 
12 

18 

1 2 
7 
18 
12 
15 
12 

12 

1 11 
1 11 
9 
12 
4 6 
15 
9 
12 



6 
6 
6 




6 
6 




TothePnblie. 



d: s. 



18 

6 

11 

Q 



6 19 

15 

11 16 

12 11 

16 

13 7 
10 

16 

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10 

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16 

1 
16 
16 

2 2 
2 2 
12 
16 

6 

1 
12 
16 



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LISTS OP THE ANIMALS IN THE SOCIETY'S GARDENS. 

List of Vertebrated Animals Living in the Gardens of the Zoological 

Society of London. 8vo. 1862. Price Is. Gd. 

List of Vertebrated Animals Living in the Gardens of the Zoological 

Society of London. (Second Edition.) 8vo. 1863. 

Price Is. 6d. 
List of Vertebrated Animals Living in the Gardens of the Zoological 

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Society of London. (Pourth Edition.) 8vo. 1866. Price Is. Gd. 
Kevised List of the Vertebrated Animals now or lately Living in the 

Gardens of the Zoological Society of London. 8vo. 1872. 

Price 2s. 
Revised List of the Vertebrated Animals now or lately Living in the 

Gardens of the Zoological Society of London. — Supplement, 

containing Additions received in 1872, 1873, and 1874. 8vo. 

1875. Price Is. 

List of the Vertebrated Animals now or lately Living in the Gardens 

of the Zoological Society of London. (Sixth Edition.) 8vo. 

1877. Price 3s. Gd. 

List of the Vertebrated Animals now or lately Living in the Gardens 

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1879. Price 3s. Gd. 

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of the Zoological Society of London. — First Supplement, con- 
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List of the Vertebrated Animals now or lately Living in the Gardens 

of the Zoological Society of London. (Eighth Edition.) 8vo. 

1883. Price 3s. Gd. 

Catalogue of the Library of the Zoological Society of London. 
(Fourth Edition.) 1887. In cloth. Price 4s. 



TJiese publications may be obtained at the Society's Office (3 
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E.O.) or, through any bookseller. 



PROCEEDINGS 



OF THE 



SCIENTIFIC MEETINGS 



OP THE 



ZOOLOGICAL SOCIETY OF LONDON. 



January 18, 1887. 
Prof. W. H. Flower, LL.D,, F.R.S., President, in the Chair. 

The Secretary read the following report on the additions to the 
Society's Menagerie during the month of December 1886 : — 

The total number of registered additions to the Society's Mena- 
gerie during the month of December was 89. Of these I was by 
birth, 71 by presentation, 5 by purchase, 6 by exchange, and 6 on 
deposit. The total number of departures during the same period, 
by death and removals, was 125. 

The most noticeable additions during the month were : — 

1. A young male of the true Zebra, Equus zebra, purchased 
December 1 1th, which fills a serious void in our collection of Equidse, 
no specimens of this now rare animal having been received by the 
Society since 1867. It would appear, however, from Mr. H. A. 
Brydon's recent letter in the Field ^, that this animal is not yet, as 
has been supposed, quite extinct in the Cape Colony. 

2. A young male of the larger Indian One-horned Rhinoceros 
{Rhinoceros unicornis), presented by H.H. The Maharajah of 
Cooch Behar, F.Z.S., through the kind intervention of Dr. B. 

' Mr. H. A. Brydon says : — " The true Zebra, the Equus montamis, the hippo- 
tigris of the ancients, the dhow of the Hottentots, and the wilde paard (wild 
horse) of the Cape Dutch, is purely and essentially a mountain-abiding animal. 
It inhabits the most remote and rugged ranges of the Cape Colony ; and at the 
present time, though sadly reduced in numbers and in the limits of its occur- 
rence, it may be found in the Sneewenburg, the Zwaart Euggens, the Zwartberg, 
and Winterhoek mountains, and in one or two other localities, in the Eastern 
Province. Quite recently a troop was running on the slopes of the Cockscomb, 
the highest peak (6000 feet in height) of the Winterhoek."— 2%e Field, vol. 
Ixviii. p. 816, Dec. 4, 1886. 

Proc. Zool. Soc— 1887, NoJ[.. 1 




2 MR. P. L. SCLATKR ON CHRYSOTIS BODINI. [Jan. 18, 

Simpson, and received December 25th. This is an important 
acquisition, as the only other specimen we possess of this huge 
animal is the male presented by the late Arthur Grote, Esq., F.Z.S., 
which has lately shown serious symptoms of old age. 

Mr. F. W. Styan, F.Z.S., placed on the table for exhibition a 
collection of eggs of Chinese birds, which he had made in the 
vicinity of Kinkiang and Shanghai. The collection contained 
clutches of the eggs of Cya7iopoUus cyunus, Chibia hottentotta, 
Acridotheres cristatellus, Corvus torquatus, Munia acuticauda, 
Rhynchcea capensis, Hydrophasianus chirurgus, Gallici-ex cristatus, 
Ardetta flavicoUis, Anas zonorhyncha, and Podiceps minor, 

Mr. Howard Saunders, F.Z.S., called attention to a specimen of 
the Mediterranean Black-headed Gull (Larus melanocephalus), shot 
on Breydon Water, near Great Yarmouth, on the 26th December, 
1886, and sent up for exhibition by Mr. G. Smith of that town. Mr. 
Saunders remarked that the bird was an adult in winter plumage 
{i.e. without the black nuptial hood), as indicated by the primaries 
being of a pure white, except a narrow black streak on the outer web 
of the first primary, a coloration which distinguishes the adult of this 
species from any other Gull of the Hooded group. An immature 
example of the same bird, said to have been shot near Barking 
Creek, on the lower Thames, in January 1866, was in the British 
Museum ; and there could be little doubt of the correctness of its 
history, which Mr. Saunders had given in 'The Ibis,' 1872, p. 79, 
and in the fourth edition of ' Yarrell's British Birds,' vol. iii. 
p. 605. The somewhat restricted breeding-area of L. melano- 
cephalus was known to extend from the Black Sea along the Medi- 
terranean to the south-west coast of Spain outside the Straits of 
Gibraltar. Mr. Saunders had also reason for believing that this 
species breeds on the shores of France south of the Gironde; it 
undoubtedly frequented that coast up to Bordeaux in winter, and 
MM. Marmotton and Vian had stated that an example taken at Le 
Crotoy, in Normandy, on the 28th of November, 1878, was in the col- 
lection of the former. South-westerly gales, such as prevailed in 
December, would easily bring a straggler to our shores. 

Mr. Sclater exhibited a skin of the rare Amazon Parrot, Chrysotis 
bodini of Finsch (P. Z. S. 1873, p. .569, pi. xlix.), brought by Mr. 
W. L. Sclater, F.Z.S., from British Guiana. 

The specimen had been obtained alive from a settler on the 
Amacuru River, British Guiana, by Mr. E. F. im Thurn, in October 
last, and kept for some time living at Maccasseema, his residence 
on the Pomeroon. This Parrot was stated to be known to the 
Warrau Indians of the Amacuru district as the " Toua-toua," and 
to be found wild in the mountainous district of the Upper Amacuru. 
It was considered by the Indians to be rather a rare bird, and was 
much valued for its talking proclivities. 






!lkH\Sl^ 



(D 








j^?fTf7n?s^ 



tfK tx 



%^HlStO^ 



Pi 



P,ZS. 1887. Plate II. 




-pi 



G]llWbocfera.fdlith 



29- 

Pelvic fms of CeTa-todus. 



IVeat, Nownaa-Ti 8<Co imp 



03 

r-' 

CO 

oo 






c 

a 






1 



1887.] ON THE PAIRED FINS OF CERATODUS. 3 

Mr. Tegetemier exhibited and made remarks on some heads of the 
Suraatran Rhinoceros {Rh. sumatrensis), male, female, and young, 
forwarded from Sarawak, Borneo, by Mr. Brooke-Lowe. 



Prof. Bell exiiibited a specimen of Nereis pelagica which he had 
received from his excellent correspondent Mr. R. L. Spencer of 
Guernsey, and which was remarkable for the bifid arrangement of 
tiie posterior portion of the body. He remarked that although Mr. 
Robertson, of Oxford, Dr. Horst, and himself had put on record 
Lumbrici with trifid ends, which probably were not really uncommon, 
he had not been able to find any record of a similar condition in a 
Polychsete. 

A communication was read from Messrs. H. B. Brady, F.R.S., 
AV. K. Parker, F.R.S., and T. Rupert Jones, F.R.S., containing an 
account of the Foraminifera procured on the Abrohlos Bank during 
the cruise of H.M.S. ' Plumper ' in 1857. 

This memoir will be printed in the Society's ' Transactions.' 



The following papers were read : — 

1. On the Skeleton and Affinities of the Paired Fins of 
Ceratodus, with Observations upon those of the Elasjno- 
branchii. By G. B. Howes, F.Z.S., F.L.S., Assist. Prof, 
of Zoology, Normal School of Science and Eoyal School 
of Mines, S. Kensington. 

[Received December 14, 1886.] 
(Plates I.-III.) 

Contents. V&efi 

I, Introduction 3 

II. Ou the Structure of the Ceratodus Fins in general and of the Pelvic 

Fins in particular 4 

III. On the Pectoral member of Ceratodus compared with the Pelvic one 

of the same and the Pectoral one of the Plagiostomes 11 

IV. On the proximal Postaxial Elements of the Ceratodus Pelvic Fin ... 16 
V. On the Morphology of the Axis of the Ceratodus Fin 18 

VI. On the. Homologies of the Chimieroid Fin-skeleton, as compared 

with that of Ceratodus 22 

VII. Conclusions 24 

VIII. List of Authorities referred to 24 

IX. Description of the Plates (I.-III.) 26 

I. Introduction. 

I have lately described (17, p. 277) the vertebral column of a 
Frog in connexion with which there had taken place, under disloca- 
tion of the urostyle, a process tantamount to that of reproduction of 

a lost part. While searching for literature bearing upon this subject, 

1* 



4 PROF. G. B. HOWES ON THE SKELETON AND [Jan. 18, 

there came under my notice a short paper by Traquair (26, p. 143), 
in which he describes the restoration of parts of the tail of Proto- 
pterus. Finding that he had discovered certain irregularities in the 
skeletal elements of the said restored tails, and knowing that Haswell 
had recorded (15) some irregularities of the Ceratodus paired fins, 
it occurred to me that the same determining cause might have been 
at work in the two cases — i.e., that Haswell's "branching" fins 
might perchance be " restored " ones, like Traquair's. I was soon 
undeceived ; for, apart from Haswell's paper, I have had the good 
fortune to examine one such fin, sent by him to Prof. Huxley. The 
deductions arrived at in the sequel have arisen out of a study of 
it and of the fins of five other individuals. Three of them were 
kindly lent me by my master, Prof. Huxley ; of the two which 
remain, one forms part of our teaching-collection at South Ken- 
sington ; for the loan of the other I am indebted to my Demon- . 
strator, Mr. M. F. Woodward. 

It is remarkable that Giinther, in his Monograph on Ceratodus 
(14), does not mention Traquair's discovery already alluded to. It 
is clear that that author's paper must have escaped him, as I fail to 
find note of it under either " Pisces " or " Ganoidei," as reported by 
him for the ' Zoological Record ' during both its year of publication 
and the succeeding one. 

The structural plan of the fin of Ceratodus is too familiar to merit 
detailed description here, Huxley has described (19, p. 46 e? seg.) 
its general features with exceeding care, and I shall, in accordance 
with his system, speak of the segments of the axis as " mesomeres." 

The lateral rays will be described, under the same nomenclature, 
as paratneres ; those which look dorsally when the fin is j)laced 
against the side wall of the body (anteriorly when it is held out at 
right angles thereto) I shall speak of as preaj:ial ; those which 
look ventrally under the first-named condition (posteriorly under the 
last-named) I shall describe as posiaxial. Preaxial and postaxial 
correspond to the " dorsal " and "ventral" of the Germans. As 
the basal segment of the axis differs in its essential characters from 
those which follow upon it, I shall refer to it as the proximal meso- 
mere (the " zwischen-Stuck " of Davidoff (7), the " erste Glied" of 
Schneider (23)). 

II. On the Structure of the Ceratodus Paired Fins in general 
and of the Pelvic Fins in particular. 
The majority of observations made thus far upon the fins of 
Ceratodus bear especially upon the pectoral member ; its pelvic 
representative has received less attention. DavidofF (7) and Haswell 
(15) have dealt most fully with it, the last-named author espe- 
cially as to certain "irregularities" mentioned in the Introduction. 
Fig. 1 is a faithful representation of the pelvic fin presented by 
him to Prof. Huxley ; and as it does not appear to correspond 
with any one figured in his own paper, I proceed to describe it in 
detail. 

The fin reached me cleaned and prepared, as represented in the 



1887.] PAIRED FINS OF CERATODUS. 5 

figure, and it had first to be ascertained from which side of the 
body it was derived. Its proximal mesomere (m.p., fig. 1) carries a 
large tubercle {tb.), which, as Schneider has lately pointed out, " bei 
der Brustflosse ventral, bei der Bauchflosse dorsal steht " — when the 
limb is in apposition with tiie body-wall. This process is, in all 
pelvic fins examined by me, somewhat crescent-shaped and out- 
wardly directed, its inner face being excavated. In the fin under 
discussion its outer surface was flattened ; but as its inner one sloped 
obliquely outwards, I conclude that that fin was a right-sided one. 
It is represented in the figure as seen from the dorsal aspect. 
Its axis is for the most part unequally segmented and irregular, the 
proximal mesomere being the least modified portion thereof as com- 
pared with the more normal fin. The second mesomere is greatly 
elongated, and it bears upon its postaxial border (left hand of the 
figure) a notched lobe, with which are connected five parameres. 
The two distal of these break up peripherally, and, on examining the 
individual specimen, it is hard to conjecture how far the lines of de- 
marcation between the parameres and the lobe, and between it and the 
main piece of the axis, may represent the last traces of original lines 
of separation, or the lines of cleavage of a primarily continuous sheet. 
Preaxially, the second mesomere carries five parameres ; these are 
fairly imiformly set upon it, and the distal one of the series branches in 
a true dichotomy. Interposed between the free ends of the two proxi- 
mal of these rays there is a smaller one (marked * in the figure), 
which I take to resemble those found by Davidoff (7, p. 127), occa- 
sionally lying free at the distal end of the fin. The rest of the 
skeleton is chiefly remarkable as concerns the axis ; this appears to 
be longitudinally cleft, and made up of a longer preaxial and a shorter 
postaxial piece, both of which are very irregularly segmented. All 
the parameres borne upon it, however, are simple unbranched rods, 
which difl^er from those more generally present only as regards their 
feeble segmentation. 

On examining the above-named fin with care, my attention became 
arrested by the cartilage marked r in the figure, the characters and 
relations of which are altogether exceptional. Wiedersheim has 
called attention (30) to the tact that in Protopterus the basal seg- 
ment of the axis may bear a lateral piece. To the consideration of 
this I shall return. In no regular Ceratodus fin {i. e. that bearing an 
equajly segmented axis) yet described has there been found, post- 
axially, a cartilage like the above named, attached directly to the 
basal mesomere. That element is generally held to be destitute of 
rays. Giinther has figured (14, pi. 36. fig. 4) a pelvic fin of the 
right side, which bears lateral cartilages in the above-named region ; 
but I find no mention of the fact in his text. It is to me inex- 
plicable for what reason he shoidd have failed to describe so 
remarkable a feature. I shall return, in the sequel, to the discussion 
of this fin. Haswell has figured and described (15, figs. 5, 6, 7 ') 

' His i3g. 2 is Scaid to be a representation in outline of the pectoral fin, after 
Huxley. It is unfortunate that the bouudary-Iine between the two basal 
mesomeres, indicated in the original, should have been omitted, 



6 PROF. G. B. HOWES ON THE SKELETON AND [Jan. 18, 

what I imagine may represent the cartilage in question, and that, as is 
here the case, in irregular fins. Be it, as it there exists, what it may, 
its characters in the fin figured by me are still further noteworthy. 
The entire fin-skeleton (fig. 1), with the exception of this bar and the 
proximal mesomere, is very slender and leaf-like ; the two elements 
just named (which, be it remembered, are in direct connexion) are 
relatively massive and much thicker and more powerful than the 
rest. The bar r, instead of being ellipsoidal in transverse section, 
as is invariably the case with even the most powerful parameres, 
is expanded along its free border in a manner strikingly suggestive 
of the metapterygium as it exists in many Elasmobrauchs. It is 
segmented into a main piece and two small terminal ones, and 
appears, at first sight, to represent an element of greater importance 
than an ordinary ray. 

The fact that this new element appears in "irregular" fins, taken 
in conjunction with the fact that no such structure has hitherto been 
recorded for a " regular " fin, appears at first sight to detract from its 
novelty. Before proceeding further, therefore, three questions must 
be met : — 1. How far is the fin under discussion abnormal? 2. Can 
the existence of the new element be demonstrated for a more normal 
fin ?, and 3. If so, under what structural conditions does it exist ? 
Giinther, in his original description of the Ceratodtis fin, described 
(14, p. 532) certain " slight irregularities" in the distribution of the 
rays. Bu\ley(19, p. 47), commenting upon these, remarks that 
they are " in respect of the median pieces .... constant peculiarities 
of no small importance." DavidcfP(7, p. 126) describes the stem of 
the pelvic fin as consisting of a row of pieces " deren Zahl bei den 
versehiedenen Individuen betriichtlich variirt ; " he adds — " nirgends 
fand ich eiu so unregelmassiges Verhaltniss derselben zu einander, 
wie es Giinther auf seiner Figur abbildet." Other writers have 
observed this irregularity, and the last of them (Schneider) has 
formulated the distribution of the parameres of both fins. He states 
(23, pp. 521-22), "bei der Brnstflosse sitzt dorsalwiirts am zweiten 
bis elften Gliede des Hauptstrahls, und zwar an der distalen Gelenk- 
flache, je ein Seitenstrahl. Ventralwarts sitzen am zweiten Gliede 
des Hauptstrahls hinter einander fiinfSeitenstrahlen, am dritten und 
vierten Gliede je zwei, an den folgenden einer. Bei der Bauchflosse 
tragen dieGlieder des Hauptstrahls ventralwarts je einen Seitenstrahl, 
dorsalwarts je zwei Seitenstrahlen." I have taken some pains to 
test the reliability of this very definite statement, and am in a 
position to assert with equal assurance that the only constant 
character as yet recognized is the attachment of one ray to the pre- 
axial border of each pectoral mesomere {cf. figs. 5 c&: 6). Even in so 
modified a fin as that of fig. 5, where several of tlie parameres are 
branched and two are directly confluent, this rule holds ; and in no 
regular pectoral fin yet examined has an exception to it been found. 
I give below a table of average distribution of the parameres of those 
segments dealt with by Schneider, calculated out from observations 
made upon eight pectoral and ten pelvic fins. 



1887.] 



PAIRED FINS OF CERATODUS. 



Schneider 
Observed . 



Pectoral Jin. 
Postaxial 
Seg. ii. 



5 
3-5 



111. 

2 

1-9 



IV. 

2 
1-6 



V. 

1 
1-3 



Preaxial. 
ii. 
1 
1 



Schneider . 
Observed 



Pelvic Jin. 
Postaxial. 
Seg. ii. 
. 1 
. 2-1 



Preaxial. 
ii. 
2 
2-5 



Further comment upon the pectoral member may be deferred until 
later. Concerning the ten pelvic fins examined by me, I may add 
that in eight the second mesomere bore preaxially two parameres 
(figs. 3 & 7); in a ninth three; in a tenth four. In most cases 
two postaxial rays were present (fig. /). One fin, interesting beyond 
this, bore (fig. 2, right hand, as drawn) preaxially two rays, post- 
axially four, that being a precise reversal in duplicate of the condi- 
tion observed by Schneider. In no case have I observed the distri- 
bution recorded by him. 

The parameres of all the fins alluded to were, for the most part, 
rod-like and segmented ; but in not a few instances they were 
branched or otherwise modified {cj. figs. 1, 5, 7). Reflection upon the 
facts recorded concerning them, to say the least, shakes our trust 
in the supposed regularity of their distribution. That, however, 
can no longer be asserted, in view of the truly remarkable condition 
of one pair of fins, which belonged to a fish in all respects normal 
and healthy (fig. 2). Giinther first directed attention to the sickle- 
shaped contour of the Ceratodus fin, and all subsequent observers 
are agreed as to the asymmetry of its two lobes, Schneider states 
(23, p. 521) : — " das zweite Glied des Hauptstrahls zerfallt durch 
eine Langsgrube in zwei Sliicke. Das eine Stiick behiilt die Richtung 
des Hauptstrahls, das andere Stiick divergirt mit demselben uud 
zwar bei der Biustflosse dursalwarts, bei tier Bauchflosse ventral- 
warts." And further, " Die Seitenstrahlen der dorsalen Halfte der 
einenFlosse entsprechen derjenigen der ventralen Halfte deranderen." 
A cursory glance at the pair of fins now under consideration (fig. 2 ^) is 
sufficient to show how erroneous is this deduction. That Schneider 
has accurately represented the facts for the animals at his disposal, 
I have no doubt ; but that his conclusions are incapable of a wider 
application is here proven. 

' I was at one time under the impression, from an examination of Davidoif a 
figures (7, pi. 9. figs. 6 & 7), that he had been dealing with a similar pair of 
fins ; but I am no longer in doubt. His drawing of the fin-skeleton of fig. 7 is 
not in accord with the description given, as regards the pelvis and basal 
mesomei-e. He, moreover, states emphatically (p. 127), " die Zahl der ventralen 
resp. medialen Eeihe [referring to tlie parameres] entspricht genau der Zalil 
der Gliedstiicke des Stamuies, wahrend diejenige der dorsalen resp. lateralen 
Eeihe fust genau um das Doppelte grosser ist? " 



8 PROF. G. B. HOWES ON THE SKELETON AND [Jan. 18, 

Fig. 2 represents the ventral aspect of the pair of fins afore 
named, as they lay in Hfe. They were attached to the pelvic 
cartilage (pi.) by a fibrous buffer, identical witli that described by 
Davidoff (7, p. 124). The free end of the hip-girdle terminated 
in front in a pointed extremity {processus impar of Davidoff), which, 
as already observed by Giinther (14, p. 535) and that author (7, 
p. 124), was bent towards the left side. I figure this (fig. 2 a), as 
its distortion is here much more marked than in any specimen yet 
drawn. 

According to Schneider (23, p. 521) "Die Curve, welche der 
dorsale Rand jeder Flosse beschreibt, ist verschieden von der 
Curve des ventralen Randes. Nun ist der dorsale Rand der einen 
Flosse congruent mit dem ventralen Rande der anderen." In the 
specimen here figured, the two fins were sickle-shaped ; the inner 
half of the preaxial border of the left one was straight, as repre- 
sented in the figure. It will be observed that as they lay flattened 
out, their free ends were both directed towards the animal's right 
side ; the excavated border, which imparts to the fin-lobe its sickle- 
shape, was preaxial for the right fin, postaxial for the left. When 
applied to the sides of the body, the apex of the former looked 
dorsally, that of the latter ventrally. Tlit contour of the Ceratodus 
fin is variable ; occasionally its opposite margins are symmetrical 
with respect to the axis ; but the differences in symmetry between 
these two fins more than cover those which I have observed between 
any two members at my disposal. Turning now to the supporting 
skeleton, it will be seen that the second mesomere bears, as Schneider 
has pointed out, an accessory lobe (his "anderes Stiick " referred to 
above). That, however, instead of being symmetrical, as he claims 
it to be, is, in this specimen, unsynimetrical to tlie utmost — for the 
right fin it is postaxial, for the left one preaxial. Further comment 
is needless, as the drawing which I give speaks for itself. Thus far 
the characters ol the pelvic fin, as defined by Schneider, are seen to 
be inconstant and untenable : more than that, however ; for, in that 
the preaxial lobe of the one fin corresponds almost to a degree 
(with the exception of one feature, to which I shall return) with the 
postaxial lobe of its fellow and vice versa, there are embodied in the 
two the more important differences held by him to exist between 
the pectoral and pelvic members. 

Schneider goes on to say (p. 523), " weun man die symmetrische 
Stellung der vorderen und hinteren Flosse in Betracht zieht, so 
leuchtet die Aehnlichkeit des ersteu Gliedes des Hauptstrahls mit 
Humerus und Femur des zweiten Gliedes des Hauptstrahls mit 
Ulna-Radius und Tibia- Fibula ein." I ha\e shown above that the 
characters of this "zweites Glied " are inconstant for the pelvic fin. 
Its accessory lobe is present on that side on which the parameresare 
stoutest, be it preaxial or postaxial ; and examination of the 
specimen under my hand suggests unmistakably that it has arisen as 
the result of coalescence between the second mesomere and the 
confluent bases of the two proximal parameres. The well-known 
lobe of (he pectoral member (e/. figs. 5 and 6, int.), fir;t accurately 



1887.] PAIRED FINS OF CERATODUS. 9 

described by Huxley (19, p. 49), to which Schneider likens that of 
the pelvic fin, is constant in its relationships and invariably post- 
axial. I emphatically deny that structural similarity of the second 
mesomere of the fore and hind fins suggested by him, while I desire 
to lodge a protest against the unqualified assertion that (23, p. 523) 
" das Problem der Entwickeiung von Arm und Bein, welches gegen- 
wartig so vielfach behandelt worden ist, wird dadurch .... seiner 
Losuug einen Schritt naher gefiihrt." 

The great variation here demonstrated in the relative number 
and calibre of the parameres of opposite sides of the normal pelvic 
fin at least shows that the numerical differences existing between 
them and those of the so-called irregular fin described at the outset 
are insignificant. What now of the " branching," to which attention 
was originally directed by Haswell (15, p. 7) ? In the fin furnished by 
him all the rays not indicated in the drawing (fig. 1) are simple and 
unbranched, though somewhat unusually elongated. Many of them 
are transversely segmented. The question resolves itself into this — 
Can the irregularities represented in fig. 1 as it stands be shown to 
exist in a more normal fin ? Bifurcation of the terminal portion of 
one or more parameres is no exceptional feature. Giinther (14) 
and Davidoff (7) have both described it for the pelvic fin, and I 
figure an example (fig. 7) in which it had attained a marked develop- 
ment. Fig. 5 shows that it is no new peculiarity for the pectoral 
fin also'. I have seen a dichotoiny of the pectoral paramere in one 
other case, and that in a fin in all other respects normal. The trans- 
verse segmentation of the axis of Haswell's fin (fig. 1) is not a whit 
more remarkable tliau that of fig. 7 ; while in the fin there repre- 
sented, as in the pectoral one of fig. 5, irregularities of the preaxial 
parameres existed which far exceed in abnormality (if such it may 
be termed) anything forthcoming in the first-named specimen. 
Briefly stated, Haswell's fin differs most conspicuously from that 
of the more constant type in respect to the longitudinal cleavage 
of the axis. This phenomenon has already been recorded by 
Haswell, and that in a fin which recalls the one here described 
(15, pi. 1. fig. 6). Albrecht has figured and described (Sitzungsb. 
d. koaig. preuss. Akad. Berlin, vol. xxxii. p. 545, 1886) a specimen 
ol Protopterus (P. annectens) in which the distal half of the axis of 
the left ])ectoral fin had similarly bifurcated ". 

Haswell (15, p. 8), commenting upon the " branching" process 
which he first described, asserts the belief that " it is reasonable to 

^ I found, ou examiniug this specimen minutely, that many of the parameres 
terminated in small nodules such as are represented at *. On comparison with 
the other specimens dissected by me, I am convinced that similar terminal seg- 
ments existed in two cases, but that, owing to their delicate nature, they had 
been for the most part torn away in the process of dissection. The free ends of 
the rays from which they had been thus removed presented a characteristic 
truncated appearance, identical with that represented in some of the rays so 
carefully dravyn by Davidoff (1). Putting all together, I incline to the belief 
that the terminal nodules in question are of tairly general occurrence. 

^ The deductions which he has drawn from the study of this fin appear to me 
no less unwan-antable than those of Schneider alluded to above. 



10 PROF, G. B. HOWES ON THE SKELETON AND [Jan. 18, 

regard it as an instance of atavism, and so pointing back to a pre- 
existing condition in which the fin-skeleton consisted of branching 
jointed cartilaginous elements supporting a cutaneous expansion 
considerably broader than that of tlie fin of the living Ceratodus 
forsteri." If, as therein suggested, the typical paramere has 
arisen from a confluence of branching-elements, such as exist 
to-day among some Elasmobranchs, and if it be that the meso- 
meres have been formed by the fusion of the basal ends of the 
parameres as they now stand, each mesomere would be morpho- 
logically double, and the longitvidinal cleavage of this axis would 
thereby receive an intelligible interpretation. I am doubtful as to 
the probability of such a process having been involved, but, in the 
absence of any data upon the development of the fin, I put forward 
the suggestion as a possible means of accounting for the apparent 
irregularity. In support of this conception of the origin of the 
parameres, it may be stated that their reduction in number is 
proportionate to the thickening of the fin border. Schneider says 
(23, p. 521) that "die Seitenstrablen der dorsalen Hiilfte der einen 
Flosse entsprechen derjenigen der ventralen Hiilfte der anderen." I 
find, however, that when (as in the left-hand fin of fig. 2 — right-hand 
one of the drawing) that lobe which is generally thickened remains 
thin, its supporting rays are more numerous and of smaller calibre 
than usual. When, on the other hand, as was also the case in the 
fin represented, the usually thin lobe becomes thickened, its support- 
ing rays get less numerous in proportion as they become more 
powerful. Stated otherwise, these facts go far to prove that the 
thickening of one or other of the fin- borders is mainly due to 
confluence and subsequent increase in calibre of the parameres. 
Suggestive, indeed, in view of all this is tlie occasional bifurcation 
of a linear series of postaxial parameres, such as is represented in 
fig. 7. 

Perusal of the foregoing pages will show conclusively that 
Haswell's " branching " fin is, when compared with those of a number 
of other individuals, little, if at all abnormal. There yet remains for 
consideration that cartilage {r, fig. 1) which, as I have stated, is 
connected with the proximal mesomere ; and it has now to be incjuired 
if a representative thereof is forthcoming in a more typical fin. 
After long searching I found an unmistakable representative of 
it, and that in none other than the left fin of the remarkable pair 
represented in fig. 2. Fig. 3 is a drawing of the upper third of 
the same. The postaxial lobe was supported, as has been already 
stated, by a series of delicate parameres, of which there were two 
to each of the ray-bearing mesomeres figured, with the exception 
of the first and third {cf. fig. 2). The preaxial lobe was, contrary 
to the general rule, supported by a series of larger and more powerful 
parameres ; of these there was one to each of the above-named 
segments, with the exception of the first. None of the parameres 
showed the slightest trace of branching. 

Proximally to the postaxial rays there lay the cartilage, r, of 
fig. 3. This element was relatively far smaller than was that of the 



1887.] PAIRED FINS OF CERATODUS. 11 

specimen first described (fig. ] ), and its relationships to the basal 
mesomere were the less definite of the two. It was here segmented 
into two pieces, while it was much more intimately connected with 
the two adjacent parameres than was the case in the former 
example. These two fins (figs. 1 and 3) stand alone, among those 
which I have examined, with respect to the great increase in number of 
the parameres of the postaxial lobe, and that also bears the cartilage 
now in question. If, as Haswell suggests (15, p. 8), this duphcation 
of rays is reversionary to a " a pre-existing condition in which the fin- 
skeleton consisted of branching, jointed, cartilaginous elements," 
the only conclusion which seems to me justifiable is that the appear- 
ance of this new element amounts to that of the reappearance of 
one which has been lost. Haswell has described an individual (15, 
figs. 6 and 7) in which the cartilage in question appears to have 
been present on both sides ; and it is instructive to remark that in 
both fins the rays of the postaxial series were, as with my specimens, 
the more numerous. I have already stated that in the fin described 
at the outset (fig. i), the whole skeleton was, with the exception of 
the bar in question and the basal mesomere, very slight and leaf- 
like. This simplification of structure, so suggestive of the reversion 
claimed by Haswell, is seen in the basal mesomere itself. That was 
(fig. 1, m.p.) much thinner and more flattened than is usual, and it 
bore but one processus muscularis {tb.) instead of the three described 
by Davidoff (c/. 7, pi. 8). All the foregoing facts point to the 
conclusion that the newly described cartilage exists only in fins whose 
postaxial rays remain little modified. There is, therefore, good 
reason to regard it, let its homology prove to be what it mav, as 
atavistic. It has disappeared in the normal fin, under a confluence 
of the parameres of its own side, and a consequent thickening of the 
postaxial fin-lobe. 

III. The Pectoral member of Ceratodus compared with the Pelvic 
one of the same and the Pectoral one of the Plagiostomes. 

Hasv?ell, reviewing (15, p. 5) the well-known observations and 
hypotheses of Balfour (1), Thacher (24, 25), and others, which 
led them to dissent from the interj^retations of Gegenbaur and 
Huxley, says that they, together with the facts which he brings 
forward, seem to place it beyond a doubt that the limb of Cera- 
todus, "so far from representing a primitive and generalized 
type, is, as, indeed, we should expect from various other points 
in the organization of the animal, in reality highly speciahzed, and 
to be regarded as derivable from such simple limb-skeletons as those 
of the Selachii." In this he was anticipated by Balfour (I) whom 
he quotes. Balfour wrote (p. 669), when criticizing Huxley's 
position, the leading tenets of which he supported so far as the 
identification of the chief constituents of the fin-skeleton go, " I 
should be much mere inclined to hold that the fin of Ceratodus has 
been derived from a fin like that of the Elasmobranchs, by a series 
of steps similar to those which Huxley supposes to have led to the 



12 PROF. G. B. HOWES ON THE SKELETON AND [Jail. 18, 

establishment of the Elasmobranch fin, but in exactly the reverse 
order'". 

The researches of Huxley and Balfour have proved that the 
propterygium of Gegenbaur (figs. 9 and 10, pt.) represents, 
throughout the Elasmobranch series, but one or more preaxial rays. 
It is the most variable of the three basal elements of the Shark's 
fin, and most observers are now agreed as to its morphological un- 
importance. The above-named writers are further at one in their 
estimate of the morphological value of the Elasmobranch meso- 
and metapterygia {ms., int., figs. 9 and 10). That they disagree, 
however, upon at least one vital issue is well known, and the 
balance of opinion holds to-day that the solution of the ' archi- 
pterygium ' question is to be sought in a reconciliation between 
their views. 

Huxley has described and figured (19, p. 48) the maximum 
development yet observed for the so-called propterygium of the 
Ceratodus pectoral fin. That structure cannot be definitely re- 
cognized in the pelvic fin. The determination of Huxley (19), 
Balfour (1), and v. Rautenfeld (22), which regards the axis of the 
Ceratodus fin as the mesopterygium, is too familiar to call for com- 
ment here. It must suffice to state that I accept it in the main, if 
not wholly, and assume for the present that the entire axis has the 
value which Huxley first assigned to it. 

It is at this point necessary to discuss, more fully than heretofore, 
the nature of the differences between the pectoral and pelvic fin- 
skeletons of Ceratodus. Schneider has asserted (23, p. 521) that 
" die Seitenstrahlen der dorsalen und ventralen Halfte der Flossen 
sind ungleich," also that the " Seitenstrahlen der dorsalen Halfte 
der einen Flosse entsprechen derjenigen der ventralen Halfte der 
anderen." There is an undoubted tendency towards the assumption 
of the condition which he thus formulates for Ceratodus, and it 
seems to me probable that a common determining cause may have 
led up to it and the condition realized in Frotopterus {cf. Schneider, 
p. 524) ; but the definition no longer holds invariable for the former 
animal, in view of the facts thus far adduced. I have already 
stated that the presence of one preaxial paramere in connexion 
with each mesomere is a constant character of the Ceratodus 
pectoral fin, and I turn now to the distribution of the postaxial 
rays. I have given on p. 7 the average distribution for eight 
pectoral fins examined. The minimum observed was, taking the 
mesomeres in order of succession from within outwards, 3.1.2.1, 
the maximum 4.2.2.2. In no case have I observed five rays in 
attachment with the second mesomere, as stated by Schneider. Of 
the eight specimens examined, the second mesomeres of five bore 
each three rays ; the third and fourth of seven each two ; and the 
fiftii of six each one. It is thus certain that variation in the distri- 
bution of the postaxial parameres (c/. fig. G) is, beyond doubt, far 

' Giiuther originally advanced a somewhat similar opinion (14, p. 534), 
but he conceived of the process as having gone on along lines as yet incapable 
of support. 



1887.] PAIRED FINS OF CERATODUS. 13 

less marked than with the pelvic member ; but the fact which 
stands out most clearly is that the second mesomere invariably bears 
the greatest number of these rays. They are carried (figs. 5 and 6, 
mt.) upon a special lobe of the axial cartilage (" das divergirende 
Stiick " of Schneider) already alluded to. The free border of this 
lobe slopes, in every case examined by me, gradually towards the 
proximal mesomere {m.p.), and it is, moreover, in all, marked off 
from the body of the second mesomere by a deep furrow (indicated 
in the figs, by a dotted line). 

I now proceed to discuss its homology, and having arrived at the 
conclusions to be formulated in the sequel through a comparison 
with the pectoral fin of Cestracion, I pass at once to the considera- 
tion of that. 

Gegenbaur and Huxley are both agreed that the base of the 
Cestracion fin is supported by two cartilages (fig. 10) held by them 
to represent the mesopterygium (ws.) and the metajiterygium (tni.) 
of other Selachians. Most recent writers have adopted their views 
(cf. Hubrecht and Sagemahl in Bronn's ' Klassen und Ordnungen 
des Thier-Reichs,' vol. vi. part 4, Pisces). Huxley, instituting a 
comparison (19, p. .tO) between the corresponding fins of Cestracion 
and Notidanus, regards them as representative of the transition 
stages in the shortening of the Ceraiodus-like " archipterygiuni," 
by which he concludes the typical fish-fin has arisen. Gregenbaur 
(9, p. 148) likens the Cestracion fin to that of Acanthias, and says 
•' das Propterygium fehlt gjinzlich." 

Huxley, holding further that the propterygium (preaxial ray) of 
Cestracion is removed from the shoulder-girdle, as in Ceratodus, 
ftsserts that in Scyllium (pp. 50-52) " the further shortening of the 
axis gives rise to still greater changes. The axial cartilage (nieso- 
pterygium) is relatively small ; but the enlarged postaxial cartilage 
{metupterygium) has extended upwards along the postaxial face of 
the first, until it has not only reached the articular surface of the 
pectoral arch, but furnishes a large part of the articular cavity. In 
like manner the proximal preaxial ray {propterygium) has ascended 
along the preaxial face of the axial cartilage, until it also is able to 
furnish a facet which completes the anterior part of the cup for the 
condyle of the pectoral arch." He holds therefore that the pec- 
toral fins of Notidanus, Cestracion, and Scyllium represent, in the 
order enumerated, the successive steps in the modification alluded 
to above, and he, in accordance with the statements quoted con- 
cerning the propterygium and metapterygium, relegates the two 
former fins to his category of the "unibasal " type, as distinguished 
from that of the latter animal, which he holds to typify the " tribasal" 
one predominant among the Plagiostomes {cf. table which accom- 
panies his essay). 

From an examination of the fins of two young Cestracions, I can 
state without further hesitation that the mesopterygium of the 
adult is (as Mivart has suspected, 21, p. 477) a compound of the 
pro- and mesopterygia. Fig. 9 represents one of the fins referred 
to. The animal died at the period at which the two {ms. and pt.) 



14 PROF. G. B. HOWES ON THE SKELETON AND [Jail. 18, 

were beginning to unite. The propterygium {pt.) is seen to com- 
pose fully the anterior third of the whole mass, its base being 
made up of two segments, the proximal one of which contributes 
nearly half the articular cup, entering at least as fully into the 
formation of the same as does its representative in Scyllium (cf. 
Huxley, 19, p. 48, fig. 10). In the adult fin (fig. 10) the original 
boundary line between the pro- and mesopterygia is represented by 
a groove indicated in Huxley's figure (here reproduced) by a 
dotted line. 

The pectoral fin of Oestraeion is thus shown to conform to the 
Selachian type, being identical most nearly with that as represented 
by Acanthias (cf. Gegenbaur, 9, pi. 9. fig. 4, and Mivart, 21, 
pi. 77. fig. 2). Cestracion must, on the evidence now forthcoming, 
relinquish its position in the series established by Huxley ; it must, 
to say the least, change places with Notidanus. The main articu- 
lation of the fin of ihe last-named fish is established, as is well known, 
through the agency of the mesopterygium. Gegenbaur originally de- 
scribed a basal preaxial bar in Notidanus, and he homologizes it (9, 
p. 140) with his propterygium. This enters, if anything, more fully 
into the articulation with the shoulder-girdle than does its representative 
in Cestracion. There is connected with its distal end, in Hexachus, 
a smaller piece (c/'. Gegenbaur, op. cit. pi. 9. fig. 1) which appears 
to represent the second segment of Cestracion, reduced, as an 
outcome of the great expansion of the front border of the meso- 
pterygium. Mivart has suggested (21, p. 444) that the mesoptery- 
gium represents a coalescence of the pro- and mesopterygia ; but I 
am inclined, upon careful examination of the specimen under my 
hand, to dissent from that view. 

The metapterygium of Notidanus enters into a feeble but definite 
connexion with the pectoral arch, such as is not the case in Cestra- 
cion. That the fins of these two genera differ from those of some 
of the Sharks is indisputable, but they do so to an insignificant 
degree, incapable in itself of supporting the " unibasal " type ; that, 
in face of the facts here adduced ', rests upon an insufficient basis. 

Returning now to Ceratodus ; the lobe which, in the pectoral fin, 
carries the 3-5 j)roximal postaxial parameres (figs. 5 and 6 mt.) is, 
as has been stated previously, marked off from the adjoining meso- 
mere by a deep furrow. 

Giinther observes (14, p. 532) that the conjoined mass shows, 
in "horizontal section," lines of "the former divisions" into what 
he holds to correspond to the " three carpals " (pterygia) of most 
Plagiostonies. This has been denied by Huxley (19, p. 47). Set- 
ting aside this difference for the moment, I desire to call attention 
to the similarity of the furrow described above to that which 

^ Wlien, moreover, it is considered that in the pectoral fin of Polypfenis, 
which Huxley relegates to the " tribasal '" category, the mesopterygium is (as 
Gegenbaur pointed out, 10, p. 139) excluded from articulation with the Umb- 
girdle, the statement that (p. 55) ''the mesopterygium is the proximal piece of 
the axial skeleton, which constantly retains its primary articulation with the 
pectoral arch," must needs be modified. 



1887.] PAIRED FINS OF CERATODUS. 15 

marks off the propterygium from the mesopterygium in the adult 
Cestracion. On a comparison of the two, I submit, with some 
degree of confidence, the opinion that the postaxial lobe of the 
second mesomere of Ceratodus (fisis. 5 and 6, mt.) is the homologue 
of the Elasmobranch metapterygium. Comparison of that lobe and 
its attached rays (fig. 6) with the metapterygium of Cestracion 
and its rays (fig. 9) reveals a striking similarity, even in detail, 
betvpeen the two. Did the metapterygium of the Shark unite, 
as does its propterygium, with the axial plate ms., it would be 
difficult indeed to find a distinction between the first named and 
that which, in Ceratodus, I claim as its homologue. 

The homology which I here seek to establish bears out, with 
certain modifications, Balfour's view cited that " the fin of Ceratodus 
has been derived from a fin like that of the Elasmobranchs." That 
observer first recognized (1, p. 668) that the metapterygium (his 
basipterygiutn) is morphologically the most important and, phyloge- 
netically, the most primitive of the basal elements ; while he suspected 
(ibid.), hue did not demonstrate the fact, that that structure is formed 
by the coalescence of rays. Huxley had already asserted this belief, 
in dealing with the metapterygium oi Notidanus (19, p. 50), which 
he regarded as being " formed by the coalescence of the axial ends 
of the postaxial rays" (presumably on the shortening of the fin 
axis). Dohrn has recently substantiated the deductions of Thacher, 
Mivart, and Balfour under this head, in having found that the meta- 
pterygium is (8, p. 174), in both pectoral and pelvic fins of the 
Shark, like the basal bar of the median fins, made up of "unpaare 
Knorpelstrahlen, die anfanglich oder jede Verbindung mit anderen 
Skelettelementen bleiben." He reiterates the statement on p. 182 
in the words " was als Basipterygium beschrieben ist, stellt nur die 
verschmolzenen, wei sehr uah an einander leigenden, Basen der 
Flosseustrahlen dar und existirt nicht unabhangig von diesen." 

In face of the above facts, my view demands that a primary dis- 
tinction shall be demonstrated between the second pectoral mesomere 
in Ceratodus and that lobe which I hold to represent the meta- 
pterygium. In Giinther's original specimen (fig. 8) tiie said lobe 
was not represented in that which is now known to be its typical form, 
while the rays (r) which are usually attached thereto were for the 
most part independent. The proximal two of these appear to have 
been somewhat smaller than usual, but it is highly interesting to 
note that the two distal ones were uniting at their bases to form 
a plate-like structure {mt.) which showed no signs of confluence 
with the adjacent mesomere. This plate corresponds in its mode 
of origin with the metapterygium (basipterygium of Balfour), as 
defined by the above-named authors, and, in its relationships to the 
rest of the fin-skeleton, with the lobe now under consideration. 
I regard its condition as there represented to be indicative of the 
primary independence which my interpretation necessitates. 

Giinther goes on to say (p. 532) that he found " lines of the 
former divisions" of the second mesomere of this specimen pre- 
served, in the shape of tracts of white connective tissue. Huxley 



16 PROF. G. H. HOWES ON THE SKELETON AND [Jail. 18, 

denies the existence of these, as has previously been stated ; but let 
them be present or not, it is certain, should Giinther's observation 
hold good, that they cannot indicate the original lines of separation 
between pro-, meso-, and metapterygia, as now understood. 

IV. On the proximal Postaxial Elements of the Ceratodus Pelvic Fin. 

The cartilage which I have already described (figs. 1 and 3) as 
directly connected with the postaxial border of the basal mesomere 
of the Ceratodus pelvic fin is ray-like, but relatively powerful, in 
one of the two specimens (fig. 1). In the other (fig. 3) it is alto- 
gether smaller and segmented into but two pieces, instead of into 
three, as in the former specimen. While it here meets the distal 
end of the proximal mesomere, it is much more intimately con- 
nected with the second piece of the axis than in the former speci- 
men ; but on the supposition that the cartilage is homologous in 
both fins, its condition in fig. 3 is precisely that which would result 
from a further reduction of that of fig. 1, such as there is good 
ground to believe, for reasons previously alleged, has actually gone 
on. In the second specimen the cartilage in question is further 
interesting, in that it bears one and is in close connexion with a 
second of the proximal parameres. 

In the specimen figured by Giinther (14, pi. 36. fig. 4) already 
referred to (p. 5), the proximal piece of the axis bears two carti- 
lages. Fig. 4 is a reproduction of his drawing. The distal carti- 
lage is ray-like, and stands related to the base of the proximal 
mesomere as does an ordinary postaxial paramere to the corre- 
sponding border of a typical mesomere. The proximal cartilage 
appears to have been free of the basal piece altogether. It is, as 
shown in the figure, plate-like, and I have little doubt but that it was 
formed by the confluence of the basal ends of at least the two rays 
whicb it carries. These skeletal elements, as they stand in Giinther's 
specimen, combine the cliaracters of those of the two described by 
me (figs. I & 3). The postaxial parameres are, as in my specimens, 
much simpler than usual, and the whole series of lateral rays are in 
his fin more uniformly distributed than in general. The basal 
plate is (fig. 4, mt.), like the corresponding bar of fig, 1, in near 
relationship with the proximal mesomere, although but loosely 
connected therewith ; while it agrees with the corresponding element 
of fig. 3 in giving attachment to a couple of rays. Comparing the 
proximal postaxial elements of my two specimens and Giinther's 
figure with the corresponding region of the pectoral fin-skeleton, and 
reflecting that the typical metapterygium is formed by a confluence 
of the basal ends of the rays of that region, I incline to the belief 
that the vestiges in question represent that lobe of the fore limb 
which I claim as the metapterygium, together with its associated 
rays. 

Should the cartilages now under discussion have the morpholo- 
gical value which I am seeking to establish for them as probable, 
the well-known views of Gegenbaur (10, II), and Huxley (19) will 



1887.] PAIRED FINS OF CERATODUS. 17 

receive refutation, proportionate to the support furnished for those 
more especially of Balfour, Haswell, and Dohrn already cited. 

As stated previously, the cartilage r of fig. 1 is ray-like, but 
stouter and more powerful than that of any ordinary paramere. 

In seeking light on this question, one naturally turns to Poly- 
pterus, the aifinities between which and the Dipnoi, originally 
pointed out by Huxley (18), have nowhere been denied. The 
Polypterus pectoral fin is, as is well known, supported upon three 
basal elements. The mesopterygium (fig. 11, ms.) is held by 
all to represent that of the Plagiostomes, and no oue has yet 
challenged Gegenbaur's determination (9, p. 148) of the homology 
between the elongated postaxial bar (jnt.) of this fish and the 
metapterygium of the Plagiostomes and Chimceroids. Huxley says 
of this fin (19, p. 53) that "the Scyllium type is essentially 
preserved." Comparison of the Polypterus pectoral fin (fig. 11) 
with the pelvic fin of Ceratodus represented in fig. 1 would appear 
at first sight to suggest a homology between the basal postaxial bar 
(r) of the latter and the metapterygial bar (mt.) of the former. 
If this be justified, it would further appear, accepting the homology 
of the metapterygium of Polypterus with that of the Elasmo- 
branchii, that the two fins might have been derived along a line of 
modification characterized by the assumption on the part of the 
metapterygium of a ray-like character, and by the subsequent 
elongation of the mesopterygial plate {ms.). The probable truth 
of the latter assertion seems to me very great indeed. The meso- 
pterygium is, in Pohjpterus (fig. 11, ms.), already elongated beyond 
the limits met with elsewhere, displacing in the process the mar- 
ginal rays. Continue that elongation, and there could only result a 
Ceratodus-\\ke product. As concerns the former supposition, how- 
ever, comparison of the fin-skeletons represented in figs. 1, 3, & 4 is 
sufl3cient in itself to show that the proximal postaxial ray of fig. 1 most 
probably represents the distal one of those related to the proximal 
mesomere of fig. 4. Comparison of the latter (fig. 4) with the 
proximal end of the pectoral fin cf the same side of the same animal 
(fig. 8) shows unmistakably that in the plate-like structure result- 
ing from the fusion of the basal ends of the two proximal para- 
meres we have to deal with the last trace of the metapterygium, 
defining that, as must now be done, as a product of the confluence 
of the inner ends of the proximal postaxial rays, the distal ray being, 
from the nature of its relations therewith, one of the same series. 

Consideration of the above facts renders the homology of the sup- 
posed metapterygium of Polypterus somewhat doubtful. Gegen- 
baur, when pointing to the same, realized the similarity between both 
pro- and metapterygia so-called by him (fig. \\,pt. andrn^.) and the 
marginal rays'. He at first suggested (10, p. 139) the possibiUty 
that the exclusion of the mesopterygium from connexion with the 

^ The difficulty of interpretation of the supposed propterygium is greatly 
increased by the presence of the cartilage marked * in fig. 11, — by no means the 
least puzzling element in this fin. As wiU be seen, it is grafted upon the ante- 
rior border of the propterygial rod ; from it, however, it is perfectly distinct in 

Proc. Zool. Soc— 1887, No. II. 2 



\8 PROF. G. B. HOWES ON THE SKELETON AND [Jan. 18, 

shoulder-girdle may have been due to a displacement of the same 
by two rays. Should this be so, the metapterygiura must there 
have disappeared, as from the Ceratodus pelvic fin, under the 
corresponding enlargement of the mesopterygial plate. The only 
alternative view possible is that the metapterygium does repre- 
sent that of the Elasmobranchs. If this be so, comparison of the 
pectoral fin of Polypterus (fig. 11) with that of the Plagiostomes, as 
represented in Scyllium, where the mesopterygiuni is relatively small, 
would seem to show that the loss o^ connexion between the meta- 
pterygium and its ravs has been to no small extent due to a 
displacement of the latter by the elongation of the ex[)anding 
mpsopterygium, no less than by the simplification in structure of the 
metapterygium itself. The last step in the former process would 
appear, indeed, to be retained in the living Polypterus (** fig. 11). 
In the absence of embryological data further discussion of this 
difficulty would be fruitless. It is much more pertinent to observe 
that in both Ceratodus and Polypterus the initial step in the modi- 
fication has been, in any case, one of elongation of the mesoptery- 
gium, and evidence has been adduced to show that in the Dipnoi (if 
not in Polypterus also) the metapterygium has been thereupon 
reduced and finally suppressed. The only traces of either it or 
its rays yet recorded in the Ceratodus pelvic member are forthcoming 
in fins whose postaxial parameres are more numerous and less 
specialized than is generally the case. If this simplification in 
structure of the most specialized portion of the pelvic fin-skeleton 
is, as I have attempted to show, reversionary to a condition which 
has been lost, the characters of those elements which reappear 
under the simplification, w^hen compared with those to which they 
most nearly correspond in the pectoral fin, go far towards bearing 
out the presumed origin of the Ceratodus fin from a primarily 
expanded predecessor. 

V. On the Morphology of the Axis of the Ceratodus Fin. 

The entire axis of the Ceratodus fin is held by Huxley (19) 
Balfour' (1), and v. Rautenfeld (22) to represent an elongated 

the two specimeus examined by me. It has been figured by WiedertLeim (29, 
p. 199), but I have been uuable to find a description of it. I think it not 
improbable that it may haye been derived from the mesopterygium, the closely 
related lower anterior end of which may (as A^'iedersheim has shown) insert 
itself between the supposed propterygium and the marginal rays. In the 
absence of embryological data further discussion of it would be useless. 

* Baur has recently called attention (3, p. 6) to the fact that Gervais has 
priority over Humphrey in the enunciation of the hypothesis that the paired 
fins are dismembered portions of a lateral fold. Geryais writes (12) : — " Si Ton 
considere que les rayons des nageoires impaires des poissons out une analogic 
incontestable avec ceux dont la reunion formes les nageoires paires des memes 
animaux, c'est a dire leurs membres veritables, on est naturellement conduit a 
se demander s'ils ue seraient les homologues de ces derniers, et si Fetat d'isole- 
ment dans lequel ils restent les lines par rapport aux autres, ne resulterait pas 
de ce que chacun d'eux ne conserve pas completement ses rapports avec celui 
des segments osteodesniiques dont il est tributuire. Alors on pourrait les 



1887.] PAIRED FINS OF CERATODUS. 19 

mesopterygium, and by Gegenbaur (11) to represent the meta- 
pterygium. The points of difference between the two sets of obser- 
vers are so well known that recapitulation of thera would be super- 
fluous here. I incline most nearly to Huxley's view, and hold that 
the axis represents mainly, if not wholly, the mesopterygium of 
the Sharks ; but I regard its condition in the latter as typical of 
its earlier and more primitive state. I moreover think it not 
unlikely that the short-lobed fin of the Crossopterygidce will prove 
to be of an earlier type of structure than that of the elongated one 
of the living Dipnoi. Extended observations along the lines already 
laid down by Traquair, in his Monograph on Tristichopterus (27), 
are greatly to be desired. That the elements described by him are 
probably, and that those described by Goldfuss (13) and Kner (20) 
in Xenacanthus are certainly, homologous with those of the axis of 
the Ceratodus fin, I fully believe, and we have here the founda- 
tion of a line of study which must soon yield fruitful results \ 
Wiedersheim is the only worker who, to my knowledge, has 
offered an opinion upon the last named. He says (2D p. 195), 
speaking of the Ceratodus fin, " dass dieser Organisationsplaa 
der Brustflosse auch bei untergegangenen Fischgeschlechtern eine 
Eolle gespielt haben muss, steht unzweifelhaft fest und ich mochte 
dabei nur an den aus der Permformation stamraenden Xenacanthus 
decheni erinnern." 

There can no longer be much doubt that the confluence so 
frequently seen between one or more rays and the mesopterygium of 
the Elasmobranchs represents the last trace of the process by which 
that structure is formed {cf. Dohrn, cited on p. 15). This granted, 
it becomes a question as to how far the axis of the Ceratodus fin, 
as here defined, represents a further extension of this fusion of pri- 
marily parallel rays or an elongation of the mesopterygial plate, as it 
exists in the Sliarks. The fact that irregularity in distribution of the 
parameres is generally accompanied by that of the segmentation of 
the axis in Ceratodus {cf. especially figs. 1, 5, & 7), shows that 
there is an intimate connexion between the two ; and this is the 
more obvious on reflection that Davidoff has shown (7, p. 145) that 
the segmentation of the axis does not stand in constant relationship 
to the muscular attachments^. It is moreover inconceivable, if 

regarder comme autant cle rayons membraux rest^s libres, et ils seraient lea 
homologues de ceiix qui, par leur asservation, donnent naissance aux membres 
propremeut dits sur d'autres points du corps." 

1 I look with great satisfaction upon the work now being done in this direc- 
tion by Smith Woodward (cf. P. Z. S. 1886). It is time that some such check 
should be kept upon the deductions of the embryologist {cf. Baur, Morph. Jahrb. 
vol. viii. p. 453). 

'■' He writes, " Da wir auch an der Muskulatur eine den einzelnen Segmentea 
der Stammreilie entsprechende Gliederung fanden, zo gewinnt diese Ansichfc 
an Walirsbeinlichkeit, obwohl immerhin iiocli einzuwenden ist, dass die 
physiologische Bedeutuiig dieser Gliederung eine nur ausserst minimale sein 
kann, dass ferner auch die Zwischensehnen der Stammuskulatur in gar keiner 
naheren Beziehung zuden Segmenten der Stammenreihe steheu." I could find 
no difference between the muscles of the fin represented in fig. 6 and those 
described by him. 

2* 



20 PROF. G. B. HOWES ON THE SKELETON AND [Jail. 18, 

the Ceratodus fin has arisen as an elongation of a primarily expanded 
predecessor, that such a fusion of the approximated ends of the rays 
could have resulted from that simple process. I am of oj)inion that 
the distal portion of the mesopterygium of Ceratodus has arisen on an 
elongation of a pre-existing plate, by a process such as is seen at its 
earliest phase in Pohjpterus. 

A difficulty, however, arises with respect to the basal mesomere 
of Ceratodus, which, if it represents tlie 2)roximal end of the meso- 
pterygium, differs from that of all other fishes in forming (in the 
pectoral fin, at any rate) the sole support for the base of the fin. 
Gegenbaur, who has paid considerable attention to this matter, at 
first acquiesced (10) in Huxley's belief in the close relationship 
between Poh/pterus and the Dipnoi. Commenting upon the pectoral 
fin of the former animal, he writes (p. 138) " ausser den Selachiern 
bei denen die zweieilige Form des Archipterygium in die einzeilige 
iibergeht, besitzt vielleicht nur noch Polypterus unter den lebenden 
Ganoiden das primare Archipterygium im Flossenskelete." He goes 
on to advance the view that the fin of Polypterus represents a 
shortened-up derivative of the Ceratodus type ; but finally he reverts 
to his original position, holding, chiefly on account of the loss of 
connexion between the mesopterygium and shoulder-girdle, and of 
the great structural difference between the pectoral and pelvic fins, 
that (p. 140)"demnach kann ich das genannte Skelet von Po/?//)^erKs 
uicht uumittelbar auf das primare Archipterygium beziehen, sonderu 
leite es, wie jenes der anderen lebenden Ganoiden, von der secund- 
aren, nur eine Reihe von Radieu besitzenden Form ab." 

The pelvic fin of Ceratodus appears, at first sight, to be ex- 
ceptional in the possession of a well-developed mesopterygium. 
Davidoff has brought forward good evidence to show that the 
element hitherto regarded among Ganoids and Teleostei as the 
pelvic girdle is (5. p. 125, and 6. p. 433) homologous with the 
basal piece of the Ceratodus fin, and he terms it the basal segment 
of the metapterygium. He has shown good reason for believing that 
the true pelvis is seen for the last time among the Osteichthyes in 
Polypterus, where it is represented by two or three vestigial carti- 
lages (c/. (i. p. 4C2, pi. 21, and Wiedersheim, 28) lying imme- 
diately in front of the applied ends of the basal pterygia of o])posite 
sides. Upon careful consideration, I am disposed to accept his 
interpretation as it applies to the Ganoids and Teleostei, but I am 
more dubious about it as applying to the Dipnoi. 

Examination of either of the paired fins of Ceratodus in relation 
to the limb-girdle appears, at first sight, to favour Huxley's 
view that the whole fin-axis answers to the Selachian meso- 
pterygium. If this be so, that element must, in elongating, have 
carried down with it the metapterygium, and the propterygium if 
present. 

Balfour, criticizing Huxley's view that the basal mesomere is the 
proximal piece of the axial skeleton of the limb of Ceratodus, says 
(1, p. 669), "The entirely secondary character of the mesoptery- 
gium and its total absence in the young embryo Scyllium appear to 



1887.] PAIRED FINS OF CERATODUS. 21 

me as conclusive against Huxley's view, as is the character of the 
embryonic fin against that of Gegenbaur." 

Examination of the pectoral fin of Ceratodus shows that the 
eleiiients which I hold to represent the metapterygium and its rays 
(figs. 5, 6, & 8, mt. & r.) are related to the postaxial border of the 
second mesomere. There is at most a bare suggestion of a relation- 
ship to the proximal mesomere (m.p.). In the hind limb this is 
otherwise, for those parts of the skeleton which most nearly repeat the 
characters of the presumed metapterygium of the fore limb are 
unmistakably connected (figs. 1 & 4) with the proximal mesomere. 
If they really represent the metapterygium and its rays as they occur 
among the lower fishes, it is, I think, not unlikely, p tting together 
these facts and those recorded by Davidoff, that while the metaptery- 
gium has for the most part disappeared, the proximal mesomere may, 
after all, turn out to represent the proximal end of that structure as 
defined by Huxley, early differentiated and segmented off. 

The above suggestion, should it be substantiated, would explain 
the fact that the proximal mesomere of Ceratodus is the only con- 
stituent of the fin-axis whose characters are constant. It would 
simplify our conceptions of the fins of the Ganoids and Dipnoi, and 
bring into harmony the supposed divergent modifications of the 
fins of opposite extremities ; while it would show the pelvic member 
to be, on the whole, less modified than is usually thought. I am 
disposed to think, moreover, that it receives support from the 
absence of preaxial rays in connexion with the basal mesomere 
of Ceratodus; from the complete exclusion of the mesopterygium 
from connexion with the shoulder-girdle in Polypterus ; and from 
the condition of the pelvic fin of that animal, already alluded 
to, no less than from the marked tendency towards an increased 
development of the proximal end of the pectoral metapterygium 
among the living Ganoids, 

Still more suggestive is the condition of the basal elements of 
a Protopterus pectoral fin represented in fig. 8 a. Wiedeisheim has 
(as I have already mentioned, p. 5) shown that the proximal piece 
of the pectoral fin-skeleton of this animal bears ray-like elements. 
He describes a smaller distinct ventral (postaxial) one and a larger 
dorsal (preaxial) one, which is confluent with the main piece (proximal 
mesomere as compared with Ceratodus). I have examined two 
specimens ; in one of them the latter is much smaller than in his 
example, while in the other (fig. 8 a) there is no trace of it. I can 
only conclude therefore that it is a lobe of the basal mesomere, 
variable in character. Not so with the former ; that is in both 
perfectly distinct, being separated from the basal mesomere by a 
fibrous tract, such as subdivides any two segments from each other. 
In that specimen which was destitute of the preaxial process (fig. 8 a) 
its characters are still further noteworthy. It is elongated and 
shows traces of subdivision into two pieces, the basal one of which 
is swollen and enlarged in common with the proximal mesomere 
{m.p.), and from that it appears most clearly to have been derived. 
The second segment of the axis is in relation with both the proximal 



22 PROF. G. B. HOWES ON THE SKELETON AND [Jan. 18, 

mesomere and the swollen bafsal piece referred to ; and the whole 
condition of the parts is such as would have resulted did the proximal 
mesomere and its related lateral elements represent a shortened-up 
tnetapterygium. 

Beyond this, the pelvic member of Geratodus differs most con- 
spicuously from that of the Plagiostomes and Osteichthyes in the 
presence of an elongated mesopterygium. Balfour first showed 
(1, pp. 666-7) that the development of the pelvic fin of the Shark 
is arrested at a comparatively early stage. He and subsequent 
writers regard the enlarged preaxial ray, which Huxley would hold 
to represent the mesopterygium, as the propterygium. He has also 
called attention to the fact that the mesopterygium is not there re- 
presented ; and the only anticipation of that structure forthcoming 
among the Plagiostomes, known to me, is the comparatively insig- 
nificant one described by Haswell (16. p. 23, pi. i. fig. 3) for 
Heptanchus indicus. 

VI. On the Homologies of the Chimceroid Fin-skeleton, as compared 
with that o/Ceratodus. 

Huxley, discussing the morphology of the Chimseroid pectoral fin, 
insists (19, pp. .52-53) upon the close relationship which it bears to 
that of Geratodus. The former fin is, as is well known, supported 
upon two basal elements, botli of which are in intimate connexion 
with the pectoral girdle. The postaxial of these is held by all to 
represent the metapterygium. 

As to the preaxial cartilages: — Huxley (19, pp. 52-53), whose 
view demands that the mesopterygium " constantly retains its primary 
articulation with the pectoral arch," completely reverses Gegenbaur's 
determination (9, p. 145), and regards the smaller basal one as 
the mesopterygium, and the larger ray-like distal one which it bears 
as the propterygium. Mivart, on the other hand, insists (21, 
p. 478) on the absence of the mesopterygium, and regards both pre- 
axial elements as homologous with the propterygium. Comparison 
of the fin of Ghimcera with that of the Selachians, as represented 
in Hexanchus, appears to me to warrant his view. 

It is interesting, here, to recall Huxley's remarks upon the 
metapterygium, when dealing with Chimcera. Having asserted the 
belief that the metapterygium of Notidanus is "formed by the 
coalescence of the axial ends of the postaxial rays," he goes on to 
say (p. 53), " the metapterygial cartilage cannot, in Scyllium, at 
the same time represent coalesced postaxial rays, as the analogy of 
Notidanus would suggest, and the second joint of the axial skeleton 
as the analogy oi Ghimcera .... indicates." Hid the mesopterygium 
exist in Chimcera in the form so constant among the Plagiostomes 
— that of a fusion of the basal ends of the rays interposed between 
the pro- and metapterygia — he could, in comparing the Chimeeroid 
and Geratodus fins, only have come to the conclusion formed by 
me (p. 15), in describing the second pectoral mesomere of the latter. 

The pelvic fin of the Chimseroid is in an exceptionally interesting 



1887.] PAIRED FINS OF CERATODUS. 23 

condition. Its postaxial border is supported by a cartilage, admitted 
by all to represent the metapterygium (fig. 12, int.). This appears 
to be produced out into a preaxial lobe, which is regarded by Davidoff 
(4. p. 470, pi. 28. fig. 3, and pi. 29. fig. 18), who last described 
it, as consisting of a single piece answering to the propterygium. 

It also recalls most closely that lobe from which Balfour held (1, 
p. 667) that both pro- and mesopterygia are derived. In a young 
Chimseroid pelvic fin examined by me (fig. 12), the lobe in question is 
seen to be formed by the fusion of three preaxial rays, and careful 
examination has shown that the last traces of an original separation 
between it and the metapterygium (indicated in the drawing by a 
dotted line) exist. Did that persist, the fin would correspond in all 
essential respects with the^'pectoral member, as I have defined it ; and 
I hold that this preaxial lobe is neither more nor less than the pro- 
pterygium '. Mivart comments (21, p. 465) upon the "close 
resemblance " between the pectoral and pelvic fins of the Chimseroids. 
Comparing the pelvic fin of these animals {Callorhynchus) with the 
pectoral ones o\ Acanthias aaA Scymnus, he concludes (p. 456) that 
the basal cartilage represents, in the former, all three pterygia fused 
into one. The considerations put forward above, taken together 
with the fact that the mesopterygium never appears in the Plagio- 
stome's pelvic fin, beyond the insignificant degree observed by 
Haswell, appear to me to negative this view. 

The facts now under notice suggest, but do not prove, that the 
mesopterygium is never represented at all in the Chimseroids ; and 
that with respect to that feature those fishes stand on a lower platform 
than do the living Plagiostomes. Moreover, if the preaxial cartilages 
of their pectoral member represent the propterygium, as I believe, 
an absolute structural identity is proven between the pectoral and 
pelvic fins of the group. Both would appear to have been derived 
from the fins of an ancestor in which the mesopterygium was not 
differentiated ; and if so, that element must have been of compara- 
tively late origin. 

Davidoff has pointed to the existence of structural similarities 
between the hip-girdles of Chimeera and Cerutodus (7, pp. 142-3); 
and if the magnificent array of structural affinities between the two, 
so successfully demonstrated by Huxley (19), have the weight which 
he assigns to them, I think it more than probable, if, as I have 
suggested, the basal mesomere of Ceratodus is a derivative of the 
metapterygium, that the paired fins of the Dipnoi may have arisen, 
side by side with those of the Plagiostomes, from some such form 
as is to-day represented by Chimeera — the fusion of the rays to form 
the mesopterygium having gone on independently, the intercalation 
of that structure between the applied bases of the pro- and meta- 
pterygia, so characteristic of the Plagiostomes, having been a com- 
paratively late process. 

1 The free ray represented at * in fig. 10 has been described by Davidoflf {op. 
cit. p. 471). Tlie .spur-like outgrowth of the same, which I think may not im- 
probably represent the coalesced vestige of a second similar one, was not 
present in his specimen. 



24 PROF. G. B. HOWES ON THE SKELETON AND [Jan, 18, 

If the above suggestion should prove to have weight, the condition 
of the basal parts of the Polypterus fin, in which the mesopterygium 
is in no way in connexion with the shoulder-girdle, can only be a 
lowly one. 

It may not be inappropriate here to call attention to the conception 
lately put forward by Baur (2, p. 663j concerning the morphology 
of the cheiropterygium. He returns to Gegeubaur's first position, 
and maintains that the limb of the land-animal has been derived 
directly from the ichthyopterygium. In that case the Ceratodus fin, 
as it stands, can only represent the initial phase in a line of modifi- 
cation of the ichthyopterygium, culminating in Protopteriis (to 
include Lepidosiren. Gf. Ayers, Jenaische Zeitschr. vol. xviii. 
p. 4/9, 1885, and Schneider, op. cit.). 

DavidofF claims that the Ceratodus pelvic fin (7, p. 127) " trotz 
der Einfachheit des Gauzen, sich bedeutend komplicirter gestaltet, 
als bei den friiher bearbeiteten Fischen." He uses the words in a 
physiological sense, it is true, but that in face of his concluding 
statement that (p. 160) " das Endergebnis aber besteht darin, dass 
von der Ceratodiis-Extremitat sich diejenigeder Haie ohue Schwier- 
igkeiten ableiten lasst." This is, in my opinion, far from proven. 

VII. Conclusions. 

1. That the characters of the skeleton of the Ceratodus paired 
fins are inconstant, except for those of the preaxial parameres of the 
pectoral fin and the basal mesomere of both pectoral ar.d pelvic fins. 

2. That a metapterygium is always present in tlie fore limb, in a 
reduced condition and usually confluent with the second mesomere. 

3. That traces of what appears to represent a metapterygium are 
occasionally to be met with in the hind limb, under conditions which 
point to atavism. 

4. That the basal mesomere of the Ceratodus fin may conceivably 
have been derived from the metapterygium. 

5. That the structural features of both paired fins of the Chimse- 
roids are identical, and characterized by the absence of a meso- 
pterygium. 

6. That the paired fins of the Plagiostomes and Dipnoi have, in 
all probability, arisen independently from a type of fin most nearly 
represented by that of the living Chimseroids. 

7. Proven incidentallt/. — That the basal cartilage of the Cestracion 
pectoral fin, usually regarded as the mesopterygium, is a compound 
of the pro- and mesopterygia of other Plagiostomes. 

VIII. List of Authorities referred to. 

1. Balfour, F. M. — On the Development of the Skeleton of the 

Paired Fins of Elasmobranchii. Proc. Zool. Soc. 1881, p. 656. 

2. Baur, G. — Ueber das Archipterygium und die Entwicklung des 

Cheiropterygiums aus dem Ichthyopterygium. Zool. Anzeiger, 
vol. viii. p. 663 (1885). 



1887.] PAIRED FINS OF CERATODUS. 25 

3. Baur, G. — Historisclie Bemerkung. Month. Interuat. Journ. 

of Anat. & Hist. vol. iii. no. 1 (1886). 

4. Davidoff, M. v. — Beitrage zur vergleichd. Anat. der hinteren 

Gliedmasse der Fisclie. Morph. Jahrb. vol. v. p. 450 (18/9). 

5. Davidoff, M. v. — Continuation of the above. Morph. Jahrb. 

vol. vi. p. 433 (1880). 

6. Davidoff, M. v. — Ueber das Skelett der hinteren Gliedmasse 

der Ganoidei holostei und der physostoraen Knochenfische. 
Morph. Jahrb. vol. vi. p. 125 (1880). 

7. Davidoff, M. v. — Beitr. zur vergleichd. Anat. der hinteren 

Gliedmasse der Fische. Morph. Jahrb. vol. ix. p. 116 
(1883). 

8. DoHRN, A. — Studien zur Urgeschichte des Wirbelthierkorpers. 

IV. Mittheilung. aus der Zoolog. Stat, zu Neapel, vol. v. 
p. 102 (1884). 

9. Gegenbaur, C. — Untersuchg. zur vergleichd. Anat. der 

Wirbelth. Heft 2. Brustfosse der Fische. Leipzig, 1865. 

10. Gegenbaur, C. — Ueber das Archipterygium. Jenaisch. 

Zeitschr. vol.vii. p, 131 (1873). 

11. Gegenbaur, C. — Zur Morphologie der Gliedmaassen der 

Wirbelth. Morph. Jahrb. vol. ii. p. 396 (1876). 

12. Gervais, p.— Memoire sur la comparaison des membres chez 

les animaux vertebre's. Ann. d. Sci. Nat. se'r. 3, Zoologie. 
vol. XX. p. 21 (1853) ; also De la comp. des Membres chez les 
Animaux vertebres. Paris, 1853. 

13. GoLDFUSs. — Beitr. zur vorweltlichen Fauna des Steinkohlen 

Gebirges. Bonn, 1847. 

14. GiJNTHER, A.— Description of Ceratodus. Phil. Trans, 
vol. clxi, p. 511 (1871). 

1 5. Haswell, W. a.— On the Structure of the Paired Fins of Cera- 

todus, with Remarks on the general Theory of the Vertebrate 
Limb. Pr. Linn. Soc. New S. Wales, vol. vii. p. 2 (1882). 

16. Haswell, W. A.— Studies of the Elasmobranch Skeleton. 

Pr. Linn. Soc. New S. Wales, vol. ix. p. 71 (1884). 

17. Howes, G. B. — On some Abnormalities of the Frog's Verte- 
bral Column. Anat. Auzeiger, vol. i. p. 277 (1886). 

18. Huxley, T. H. — Preliminary Essay upon the Systematic 
Arrangement of the Fishes of the Devonian Epoch. Mem. 
Geol. Survey, dec. x. p. 1 (1861). 

19. Huxley, T. H. — On Ceratodus forsteri, with observations on 

the classification of Fishes. Proc. Zool. Soc. 1876, p. 24. 

20. Kner, R. — Ueb. Orthacanthus Dechenii, oder Xenacanthus 

dechenii. Sizb. d. Kais. Akad. d. Wiss. Wien, vol. Iv. Abth. 1, 
p. 540 (1867). 

21. MiVART, St. Geo.— Notes on the Fins of Elasmobranchs. 
Trans. Zool. Soc. vol. x. p. 439 (1879). 

22. Rautenfeld, E. v. — Morph. Untersuchg. iiber das Skelet 
der hinteren Gliedmassen von Ganoid, und Teleost. Dorpat, 
1882. ^ 

23. Schneider, A. — Ueber die Flossen der Dipnoi und die Sys- 



26 ON THE SKELETON ETC. OF CERATODOS. [Jan. 18. 

tematik von Lepidosiren und Protopterus. Zool. Anzeiger, 
vol. ix. p. 521 (1886). 

24. Thacher, J. K. — Median and paired Fins, a contribution 
to the history of vertebrate limbs. Trans. Connectic. Acad, 
vol. iii. p. 281 (1877). 

25. Thacher, J. E. Ventral fins of Ganoids. Ibid. vol. iv. 

p. 233(1878). 

26. Traquair, E. H. — On the Restoration of the Tail in Proto- 
pterus annectens. British Assoc. Reports, 1871, pt. 2, p. 143. 

27. Traquair, R. H. — On the Structure and Affinities of Tris- 

tichopterus alatus. Trans. R. Soc. Edinb. vol. xxvii. p. 383 
(1876). 

28. WiEDERSHEiM, R.— Ueber das Becken der Fische. Morph. 

Jahrb. vol. vii. p. 326 (1881). 

29. WiEDERSHEiM, R. — Lchrbuch der vergleichd. Anat. der 

Wirbelthiere. Zweite Au6age. Jena, 1886. 

30. WiEDERSHEiM, R. — Das Skelet und Nervensystem von 
Lepidosiren {Protopterus). Jenaische Zeitschr. vol. xiv. 
p. 155 (1880). 

IX. DESOEIPTION OF THE PLATES. 
Plates I.-III. 

With the exception of fig. 2, all are drawn in the same relative position. 
The postaxial (internal) border looks towards the left hand. 
Pig. 1. A " branching " pelvic fin of Ceratodus. Eight side, dorsal aspect. Nat. 
size. 

2. The two pelvic fins of a second specimen. Ventral aspect. X 1^. 
2a. The anterior portion of the pelvic girdle of the same. Nat. size. 

3. The proximal third of the left fin of fig. 2. Ventral aspect. Nat. size. 

4. Proximal third of a right pelvic fin ; indicated as seen from the ventral 

aspect, for sake of comparison with figs. 1 and 3. (After Giinther.) 
Nat. size. 

5. The left pectoral fin of fig. 2. Ventral aspect. Nat. size. 

6. Proximal portion of the corresponding fin of a fourth specimen. Ven- 

tral aspect. Nat. size. 

7. Proximal portion of the left pelvic fin of the same specimen. Ventral 

aspect. Nat. size. 

8. The proximal third of the left pectoral fin of a fifth specimen. Drawn 

to same scale as figs. 5 and 6. (After Giinther.) 
8a. Basal portion of a left pectoral fin of Protopterus. x4. 

9. The left pectoral fin of Ccdraeion fhUippi. Young J. dorsal aspect. 

Half uat. size. 

10. The basal portion of a similar fin ; same aspect. Adult. (After 

Huxley.) Reduced to the same scale as fig. 9 for sake of comparison. 

11. The left pectoral fin of a young Polypterus. Ventral aspect. x2. 

12. The left pelvic fin of a young ChiriKBra (C. moistrosa), (S . Ventral 

aspect. XH. . 
In figs. 5 and 7 the parameres not represented in full were all rod-like and 

unbranched. 
The dotted lines in figs. 9 and 12 represent lines of fusion observed ; those in 
figs. 5, 6, and 10 are inferred. 

Ecference Letters, 
cp. Base of clasper. /. Inter-articular ligament, m.p. Proximal mesomere. 
ms. Mesopterygium. mt. Metapterygium. pi. Pelvic girdle, pr. Preaxial 
fin-border, ps. Postaxial fin-border, pf. Propterygium. r. Metapterygial 
rays. 



a?- 



Ml 



Fig.l. 

(■ ■ ■ 




tPH^^ 




%V 



p.orJ.pr 



A " 



J> 



Fig. 2. 



P.Z.S.I887.P1.1V 




, -^ 






-o.o,-J I 



o^?- /aa. 



^.^^ 



■•p-»i.fir. 



Fig. 3 



i-nf.orb.pl'. 




h. I 




- ir'f CTb.pl, 









Woet , Newman. & C o luip 



P.Z.S.18S7.P1 






Fiq. 5. 
{A) 



V:,rt' -pr 



suparb pL..^ 







Fig. 9. 



Fig.n 



T. J. Pi«lt»a.ii«t. 



Figs. 5-10 CarciharodcafiroTideleti;.. Pig II, larnns. corniibica. 







^^AL H^t^ 



(■« ''>0 I 



Pi* 1! 



J.' 



id. 




/ 



c> 



.itC 



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.1 f-M—^j 


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vere.ceni (51)' V^ ^ 



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l-'ig.l4. t 



^<vd 



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v«.-l ct/u 'O; '.J^ 



ri'f <er Sv t-'r^v/RM oiirum ath. 



OaurdhaTodon rondeleui 



"v7nei,Hirwnui».3tC»Ti;" 



P.Z.S.1887.P1 VII. 



^ 



Fig. 20. 

(rial' svzej 



/ v\ 



i/^,.. 



^- 



I 






•h vLx. 



OOjlOr-^ 



I rhinen' 







Yii vui 







•r>p^/? -r • -tVi*-' 



T T . 



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iwy. 






mif cob- 



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Pt-rker &- Cowarti «i»romUtk 



opt*fT 




lob. irj^f 



Fig. 32 

(not/ sUe 




West.Newmem A Co inw- 



GsurdhaTodon rondeletii ■ 



^^^^shTmO^^ 



"^^L H\8lS^ 



pz.sise 



Fig. 24, 




V \ 




fon.t 



y 




--^^ 



Fig. 25. 




iP- /...,< 




i. h-y- 



Pi^.29 




iubr. 





•y * 


^ Fi|.2'7 




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Fig. 26 




_^ 



-m^f^^r^y^^ M 



m\'^ 



/ 






ion roiideletii (foec'^is} 



1887.] PROF. T.J. PARKER ON CARCHARODON RONDELETII. 27 



2. Notes on Carcharodon rondeletii. By T. Jeffery 
Parker, B.Sc, C.M.Z.S., Professor of Biology in tlie 
University of Otago, Ne\v Zealand. 

(Plates IV.-VIII.) 

[Eeceived November 1, 1886.] 



Contents. 



Page 
Introductory 27 

1. External Characters 28 

2. The Teeth 29 

3. The Skeleton 29 

a. Vertebral Column 30 

b. Skidl 31 

c. Skeleton of Fins 34 



Page 

4. All mentary Organs 34 

6. The Heart 35 

6. Urinogenital Organs 35 

7. The Brain 36 

Description of Plates 38 



Dr. Giinther states, in his 'Study of Fishes,' published in 1880, 
that nothing is known of the anatomy, habits, and reproduction of 
this, the most formidable of all Sharks, and that no opportunity 
should be lost in obtaining information about it. 

As no fewer than four specimens of Carcharodon rondeletii have 
been caught in the neighbourhood of Dunedin during the last six 
years, upon all of which I have been able to make some observations, 
I have decided to put these upon record, in spite of the fact that 
they are, from a variety of circumstances, detached and imperfect, 
and are very far from giving anything like a complete account of this 
very interesting Selachian. 

The following enumeration of the specimens which have come 
under my notice is given for convenience of reference. 

Specimen A. — Male, 10 ft. (3 metres) long. Caught at Moeraki, 
about 40 miles north of Dunedin, early in 1881. The viscera, 
including the heart, were removed before bringing the fish to Dunedin. 
The skeleton was prepared and is now in the Otago University 
Museum. 

Specimen B. — Female, 12 ft. 6 in. (3'8 metres) long. Caught 
in Otago Harbour early in 1885. This specimen was also evisce- 
rated, only the heart being left. Its skeleton was prepared and 
sent to the Coloninl and Indian Exhibition \ 

Specimen C. — Female, 19 ft. {5'7 metres) long. At the beginning 
of the present year two large Sharks were reported in the Lower 
Harbour, and several attempts to catch them were made by the 
local fishermen. After one or two failures (the Shark on one 
occasion having broken away with a large hook in its mouth) the 
larger of the two was caught and exhibited in Dunedin. After it 
had been on view for a few days I bought it for the museum, and 
was able to make some observations on its external anatomy, in 
spite of the advanced state of decomposition. This specimen was 
stuffed, and is now in the Otago University Museum. 

1 This specimen is now in the Natural-History Museum, South Kensington. 



28 PROF. T.J. PARKER ON CARCHARODON RONDELETII. [Jan. 18, 

Specimen D. — Female, 1/ ft. (about 5 metres) long. The second 
specimen referred to in the preceding paragraph was caught about 
a fortnight later than the first, and was also brought entire to 
Dunediu for exhibition. The funds of the museum would not allow 
of its purchase, and indeed my assistants were at the time so fully 
occupied with the preparation of the larger specimen, that it would 
have been impossible for them to undertake a second task of similar 
magnitude ; so that all I could do was to be present when, on the 
third day after capture, the viscera were removed, and to make some 
notes on those organs which were too much decomposed in the 
former specimen. 

Specimen E. — Foetus (female), 55 cm. long. For this I am 
indebted to Mr. E. P. Ramsay, F.L.S., Curator of the Australian 
Museum, Sydney. 

I may also mention that, as I am informed by my friend Prof. A. 
P. Thomas, a specimen of Carcharodon, fully 30 ft. long, was caught 
at Auckland a few mouths ago. It would seem, therefore, that in 
spite of its apparent scarcity in museums, Carcharodon rondeletii 
must be a tolerably common species in the Southern Seas. 

1. External Characters. 

As the small specimen described by Smith, in his ' Zoology of 
South Africa,' appears to be the only one of which careful measure- 
ments have been taken, it seems advisible to give a fairly complete 
series of measurements of specimen C, the largest which has come 
under my notice. 

centim. 
Total length from tip of snout to tip of upper lobe of 

tail-fin (following the curve) 577 

Total length in a straight line 552 

From tip of snout to 1st dorsal fin (anterior border) 202 
From tip of snout to anterior border of pectoral fin . . 155 
From posterior border of pectoral to anterior border 

of pelvic fin 1 52 

From posterior border of pelvic to anterior border of 

ventral (anal) fin 60*8 

From posterior border of ventral (anal) to root of 

caudal fin 40*4 

Girth immediately cephalad of 1st dorsal fin 296 

Height of first dorsal fin 45-6 

Breadth „ ,, at base 53 

Height of second dorsal fin 7 

Breadth „ „ at base 8 

Length of pectoral fin, along anterior border 103 

Breadth „ „ at base 58 

Length of pelvic fin, along outer border 30 

„ „ „ inner border 41 

Breadth „ „ at base 19 

Height of ventral (anal) fin • 11 

Breadth „ „ „ at base 9 



1887.] PROF. T. J. PARKER ON CARCHARODON RONDELETII. 29 

centim. 

From tip of snout to centre of eye 23 

„ ,, „ upper angle of nostril 22 

„ „ „ centre of mouth 30 

„ „ ,, 1st gill-slit (dorsal end) 116 

From centre of eye to spiracle 33 

Width of mouth-aperture, in a straight line 58 

Height of 1st gill-slit. 64 

Width of flattened caudal region (vide infra), 

measured with calipers 42 

Height of ditto, measured with calipers 18*5 

Height of caudal fin from tip of dorsal to tip of ventral 

lobe 146 

Height of dorsal lobe of ditto, measured along 

anterior border from root to tip 106 

Height of ventral lobe, similarly measured 95 

The skin is dark grey above, white tinged with pink below, the 
latter colour being evidently due to blood in the skin and not to the 
presence of any special pigment. The under surface of the snout 
is dark, not white and pink as in Smith's specimen. 

The snout is considerably less pointed than in Lamna, or than 
in the young specimen figured by Smith. The eye is also markedly 
smaller in proportion to the size of the head than in Lamna (cf. 
description and figures of skull, infra, p. 32 and Plates IV. and V. 
figs. 1-5 and 11). 

The form of the caudal region is remarkable, and is not adequately 
described in any of the books at my disposal, in which it is merely 
stated that the tail is provided with lateral ridges. It is more 
correct to say that the tail for a short distance in front of the 
caudal fin is strongly depressed, so much so that its width is more 
than double its height, a transverse section having the form shown 
in fig. 19, Plate VII. 

It looks very much as if this curious modification must have the 
result of providing Carcharodon (and Lamna, in which the same 
structure obtains) with a combination of vertical and horizontal tail- 
fin, the latter — the flattened region just described — being developed 
as a means of enabling the fish to rise rapidly from great depths. 

2. The Teeth. 

Only the central tooth of each row in each jaw is symmetrical, all 
the others having their long axes directed outwards. The exposed 
portion of the upper middle tooth (specimen C) is 4 cm. in height, 
and 3*7 cm. wide at the base. In the lower middle tooth these 
dimensions are respectively 3-4 (height), and 3*2 (width at base). 
In both jaws the outer surfaces of the teeth are markedly flatter 
than the inner. 

3. The Skeleton. 
The vertebral column of Carcharodon has been fully described by 



30 PROF. T. J. PARKER ON CARCHARODON RONDELETII. [Jan. 18, 

Hasse ^ and a briefer account of the entire skeleton is given by 
Haswell ^. I shall therefore confine myself to a few points which 
do not appear to have been insisted on, mentioning especially such 
as seem to be important for comparison with Lamna. 

a. The Vei-tebral Column, — In specimen A there are about 
182 vertebral centra; at the end of the tail it becomes difficult to 
count them accurately. The centra agree with Hasse's description, 
except that I do not find the difference in the disposition ol' the 
radiating lamellae of bone which that author give^ as distinguishing 
the trunk- from the tail-vertebrEe. Hasse describes and figures only 
two very thick dorsal lamellae in the caudal vertebrae, between the 
origin of the neurapophyses : in my specimens there are three or 
four comparatively thin lamellae as in the trunk-vertebrae {cf. figs. 
13 and 14, Plate Vl.). 

One point not very clearly brought out by Hasse is the extreme 
irregularity in the segmentation of the neural tube and of the 
haemal tube or ridges. These are, in the embryo, continuous 
cartilages ^, which undergo segmentation at a later stage than the 
centra, becoming divided into vertebral portions, the neur- and 
haemapophyses, and intervertebral portions, the interneural and 
interhaemal pieces, or intercalaria. The irregular way in which 
this segmentation takes place in Carcharodon is very striking, and 
is well shown in figs. 7 and 8 (Plate V.), the former representing a 
portion of the neural tube seen from above, the latter a portion of 
one of the haemal ridges seen from below, Occasionally the distal 
portion of a haemapophysis becomes segmented off, forming a rib 
(fig- 8, r). 

Another matter not touched upon by Hasse is the modification 
undergone by the vertebral column at its anterior and posterior 
extremities. Anteriorly there is no clear line of demarcation between 
skull and vertebral column. At the level of the third vertebral 
centrum (fig. 6, vert.cent. 3) the neural tube meets on each side 
with the corresponding hasmal ridge, forming a continuous lateral 
investment of cartilage over the first two centra, which are thus only 
visible from beneath. The continuous lateral cartilage thus formed 
passes insensibly into the exoccipital region of the skull, while the 
first and second centra pass into the basioccipital region, and the 
neural tube into the supraoccipital region. Thus, when the .skull 
is separated artificially from the vertebral column in such a way as 
to leave intact the great parotic processes (fig. 1, p.ot.pr), the 
plane of section passes naturally between the second and third 
vertebral bodies, and the first two centra appear to be imbedded in 
the basis cranii (fig. 3). 

It is also worthy of notice that in the first few vertebral segments 
the intercahiry pieces (fig. ti, i) are small triangular processes 

^ Das natiirliche System cler Elasmobranebier. Jena, 1879. 

^ " Studies on the Elasmobranch Skeleton," Proc. Linn. Soc. N. S. W. vol. ix. 
(1884) p. 3. 

' Baffour, ' Comparative Embryology,' vol. ii. p. 454. (Memorial edition, 
vol. iii. p. 650. 



1887.] PROF. T. J. PARKER ON CARCHARODON RONDELETII. 31 

inserted between the bases of adjacent neural arches, the latter («.«) 
forming the whole dorsal region of the neural tube ; whereas in the 
remaining greater part of the column the iutercalaria form actual 
interneural arches. 

The tail-fin contains nearly three-fourths as many vertebrae as all 
the rest of the column, the lOZth centrum being the first of the tail- 
fin (Plate VI. fig. 12, vert, cent. 107), recognizable by being the first 
to develop a haemal spine. In this and the five following vertebrae 
the haemal spine (Jice.sp) is a separate cartilage, quite distinct from 
the hsemapophyses ; in the remaining tail-vertebrae the two are 
continuous. The haemal spines gradually increase in length to the 
120th vertebra, and then undergo progressive diminution : the 
longest, in specimen A, is 10 cm. in length. 

The tail-fin is thus supported ventrally by haemal spines : the 
small portion lying dorsad of the vertebral column, on the other 
hand, has its framework constituted by a series of cartilages {ptg) 
which are evidently not neural spines but jjterygiophores ^ or carti- 
laginous fin-rays. These have no definite relation to the vertebral 
segments, one of them sometimes corresponding to a single vertebra, 
sometimes to two, and sometimes to three. 

Hasse remarks that while iutercalaria are absent in the haemal tube 
in the caudal region, they are present in the neural tube, which has 
therefore double neural arches as in the trunk region. This is true 
only as far back as the 130th vertebra (vert.cent. 130), caudad of 
which intercalary pieces are absent and the neural arches conse- 
quently single. 

In the 167th (vert.cent. 167) and following vertebrae, the neural 
and haemal arches are united with one another on each side by a verti- 
cal bridge of cartilage, so that the middle portion of each centrum is 
hidden. From the 175th vertebra to the end of the series there are 
no longer distinct neural and haemal arches, but simply an irregular 
vertical plate of cartilage, in which the last eight (?) vertebral bodies 
are imbedded. An examination of this region in the foetal specimen 
(E) shows (Plate VIII. fig. 28) that these are all perfectly' formed 
centra except the last, which is a somewhat irregular mass of bone, and 
appears to me to be a demi-vertebra ', i. e. to represent the anterior 
half of a centrum formed in the posterior moiety of the last uieso- 
blastic somite. 

In the skeletons of A and B the neural and haemal arches are 
entirely uncalcified ; in the large specimen there are small calcific 
patches on the anterior neural arches only ; from about the 4th or 
5th vertebra backwards the only calcifications are those of the centra. 

b. The Skull. — The cranium is described in detail by HaswelP, 
who gives figures of it from above and from the right side, which 
are, however, too small to show certain important details, such as 
the nerve-foramina. For this reason, and because of the desirability 

* T. J. Parker, "On the Skeleton of Eegalems argenieus," Trans. Zool. Soc. 
tol. xii. p. 24, note. 

2 Cf. " Skeleton of Begahcris," p. 22. 

' Op. cit., Journ. Linn. Soc. N. S. W. vol. ix. p. 16. 



32 PROF. T. J. PARKER ON CARCHARODON RONDELETII. [Jail. 18, 

of comparing the skull with that of the closely allied Porbeagle 
{Lamna cornubica), I give upper, under, and side views of the cranium 
of both Sharks, drawn to the same absolute length ; in the side 
views the jaws also are shown (Plates IV. and V.). 

Haswell says, " On comparing the skull of Carcharodon with a 
dried skull of Lamna cornubica, T can find little difference between 
the two." As a matter of fact the differences between the two 
crania are by no means inconsiderable, as Haswell would no doubt 
have found if his skull of Lamna had not been distorted by drying. 

The main differences are dependent upon the much greater pro- 
portional size of the rostrum and of the orbits in Lamna. In 
Carcharodon the dorso-lateral arms of the rostrum (first labial 
cartilages, W. K. Parker^) are slightly curved, with a downward 
concavity, while the median ventral arm (prenasal cartilage) has a 
strong downward convexity, so that the three bars meet at a wide 
angle. All three bars are broad at the base, but narrow considerably 
in front, and are but slightly calcified, the distal portion of the 
entire rostrum being composed wholly of cartilage. 

In Lamna^ on the contrary, all three rostral bars have a marked 
sigmoid curve, and meet with one another at very acute angles. 
They are also much longer proportionally than in Carcharodon, 
much thicker, and are covered externally with a close mosaic of 
bony matter. 

As already remarked, the eyes, and consequently the orbits, are 
proportionally much larger in Lamna than in Carcharodon ; as a 
result of this, the orbital roofs {sup. orb. pi) in Lamna are strongly 
arched both antero-posteriorly and laterally, and the infraorbital plates 
{inf.orb.pl) inclined downwards at their outer ends. The whole 
cranium also, and especially the basal plate {i. e. the basis cranii 
proper plvs the infraorbital plates), is much narrower than in 
Carcharodon {cf. figs. 3 and 4), and the parotic (p.ot.pr) and post- 
orbital {p.orb.pr) processes are less prominent. 

In the fcetal specimen (Plate VIII. figs. 24 and 25) the rostrum is 
very slender, and its ventral or prenasal bar is perforated distally by a 
foramen. The anterior fontanelle {font.) is very large, and allows 
the cerebrum to be partly seen in a view from above. The auditory 
capsules are very prominent, and show clearly the elevations for the 
semicircular canals. The supraorbital plate is hardly developed, 
and the infraorbital plate is quite narrow. 

To the outer surface of the auditory capsule of the foetus, dorsad 
of the hyomandibular facet, a small rod of cartilage (PI. VIII. fig. 26, 
spir.cart.) is attached by fibrous tissue. This appears to be the 
dorsal segment of the mandibular arch or spiracular cartilage. Unfor- 
tunately the specimen had been dissected by one of my assistants, as 
a help to the articulation of the adult skeleton, before I observed 
this cartilage, so that I was unable to make out its relations to the 
spiracle. No corresponding structure was found in the adult, but 

^ " On the Structure and Development of the Skull in Sharks and Eays," 
Trans. Zool. Soc. vol. x. (1877) p. 189. Parker and Bettany, ' Morphology of 
the Skull,' p. 35. 



1887.] PROF. T. J. PARKER ON CARCHARODON RONDELETII. 33 

in so large a specimen a small cartilage imbedded in the immense 
jaw-nmscles would be easily missed. 

The foramina in the sivull-wall have the same general disposition 
as in Lamna (cf. figs. 5 and 11, Plate V.), the main differences 
between the two being the greater proportional size of the optic 
foramen (ii.) in the latter genus, and the fact that the oculomotor 
foramen (Hi.) is on the same level as the o])tic in Carcharodon, 
wliile in Lamna it is in the same horizontal plane as the ophthalmic 
peduncle (op.ped.). In both tlie carotid foramen (cor.f.) is a short 
distance caudad of the optic. Between and below the foramina 
for the 3rd (m.) and the 5th {v.) nerves there is, in Lamna, a small 
aperture wliich does not seem to be represented in Carcharodon : 
possibly it transmits the 6th nerve. The glossopharyngeal and the 
vagus foramina (Plate IV. figs. I and 2, ix., x.) are both large, the 
latter in particular being of immense size. 

The jaws of Carcharodon (fig. 5) are chiefly remarkable for their 
great size, and esf)ecially for tlie extraordinary depth of tlie mandible. 
In Lamna (fig. 1 1) their proportional size is considerably less. 

In another closely allied genus. Alopecias, the cranium has a more 
rounded form than in Lamna, and is similarly modified in accordance 
with the great size of the eyes. The rostrum is very thin and 
delicate, and is hardly at all calcified : its ventral or prenasal bar is 
perforated at its distal end by a vertical foramen. The jaws have 
about the same proportional size as in Lamna. 

The gill-bearing arches of Carcharodon closely resemble those of 
Lamna and of Scyllium '. The hyomandibnlar and ceratohyal 
( Plate y III. fig. 27, c.hy) are large and stout, and the tongue is sup- 
ported by a flat basihyal {b.hy) having a convex anterior and an ex- 
cavated posterior border. The first branchial arch consists of aflat, 
subtriangular pharyngohyal, a stout epibranchifd, and a similar but 
longer ceratobranchial {c.br. 1) which articulates with the basi- 
hyal, there being no first hypobranchial. The next three arches 
have, in addition, a short rod-like hypobranchial segment {h.br. 2-4 ). 
Between the ventral or inner ends of tiie second hypobranchials 
(Ji.br. 2) is a small nodular basibranchial {b.br. 2), The second 
and third hypobranchials are subequal, the fourth (Ji.br. 4) is 
barely half the length of its predecessors. The fourth and fifth 
pharyngobranchials have undergone concrescence ; the fifth cerato- 
branchial {c.br. 5) is, as usual, much larger than the corresponding 
segment in the preceding arches. The last arch has no hypo- 
branchial, its ceratobranchial segment {c.br. 5) abutting against 
an elongated flattened plate {b.br. 5), rounded in front and pointed 
behind, and probably to be regarded as a fifth basibranchial. 

To the hiner face of the fifth ceratobranchial, near its dorsal end, 
a small irregular rod of cartilage is attached by fibrous tissue. Can 
this be the rudiment of a sixth branchial arch 1 

The gill-arches are but slightly calcified, even the hyomandibnlar 
and ceratohyal having only a thin crust of bony matter which does 
not extend to their extremities. 

1 W. K. Parker, op. cit. ; Gegenbaur, ' Kopfskelet der Selachier,' Ijeipzig, 1872, 
Proc. Zool. Soc— 1887, No. III. 3 



34 PROF. T. J. 1»<\.KKER ON CARCHARODON RONDELETII. [Jan, 18, 

c. The Skeleton of the Fins. — In tlie shoulder-girdle I have 
nothing to add to Haswell's description. The pectoral fin of 
specimen A exhibits a concrescence of the proximal ends of the 
mesopterjgial rays not shown in Haswell's figure. The intercalary 
pieces between the distal ends of many of the rays, referred to by 
Haswell, are evidently due to longitudinal division of the rays, one 
of which, in the specimen referred to, was distinctly bifurcated. 

The pelvic girdle and fin are not figured by Haswell : I therefore 
give a figure of those of the male specimen A (Plate V. fig. 9). 
Haswell states that the outer extremity of the pelvic cartilage {pu) 
" is produced into a process with which no fewer than six rays 
articulate." In my specimen this process is apparently represented 
by a separate cartilage («), whicli seems to be formed by the con- 
crescence of the anterior rays, and to be serially homologous with the 
propterygium of the pectoral fin. 

The first dorsal fin differs only in detail from that described and 
figured by Haswell, who says of the second dorsal and ventral (so- 
called anal) fins, that they " are very small, and consist of a few 
irregular rays without basal plates." I find, on the contrary, that 
both these fins (Plate V. fig. 10, and Plate VI. fig. 1.5) and especially 
the ventral (fig. ID) are quite typical examples of the coucrescence 
of pterygiophores (radial cartilages) to form a basipterygiura. 



4. Alimentary Organs. 

The stomach (specimen C) consists of a wide cardiac (Plate VI. 
fig. IG, card.st.) and a narrow tubular pyloric (pijl.st.) division. 
The cardiac division is about 115 cm. long and 75 cm. wide; the 
pyloric division 104 cm. long by 5 cm. wide. On the right side of 
the stom.ach, near its oesophageal end, are two hhnd pouches {x.). 

The intestine {int.) is 109 cm. long from the pylorus to the origin 
of the rectal gland, and 26 cm. in diameter. The spiral valve is 
regularly disposed, makes 48 turns, and is slightly narrower than the 
semi-diameter of the gut, so that a narrow central passage is left, as 
in Alopecias and in some specimens of Raia '. 

The rectal gland {rct.gl.) is 30 cm. long by 3"5 cm. in diameter. 
The cloaca (fig. 17) is comparatively small, and is divided by a 
horizontal fold into two chambers, an outer (c/') receiving the ovi- 
ducts (ovd.ap.), and an inner (cP) receiving the rectum (ret) and 
the urinary duct (ur.ap.). 

The liver consists of two immense lobes, which fill all the ventral 
region of the abdominal cavit}'. In specimen C the gland was too 
much decomposed'for its form and size to be made out, but in D 
(5 metres long) each lobe was about 135 cm. long, by 102 cm. wide, 
and fully 30 cm. thick. A gall-bladder is present. 

The spleen and pancreas have the usual characters ; the pancreas 
(Plate VI. fig. 16, pan.} consisting of a small ventral and a large 

' T. J. Tarker, " On the Intestinal Spiral Valves in the Genus Baia," Trana. 
Zool. Soc. vol. si. ]i. .50. 



1887.] PROF. T. J. PARKER ON CARCHARODON RONDELETII. 35 

dorsal lobe, and the spleen (spl.) being an elongated lobulated organ 
of deep red colour, attached all along the right border of the pyloric 
division of the stomach and continued on to the dorsal aspect of the 
cardiac division. 

5. The Heart. 
The heart is very large, having the following dimensions in speci- 
men C. 

centini. 

Greatest width of ventricle 19 

„ length (antero-posterior) I.t 

„ ,, of conns arteriosus 10'5 

,, width of conus arteriosus ^ 

„ ,, of auricle (moderately distended) .... 22 

Width of sinu-auricular aperture 9 

„ of auriculo-ventricuiar aperture 4' it 

Thickness of wall of ventricle (about) 3*5 

Its general structure is quite normal. Tbe sinu-auricular valves 
are obliquely right and left ; the auriculo-ventricuiar valves obliquely 
dorsal and ventral. The coronary veins o])en apparently by a single 
very large aperture situated in tbe sinus venosus immediately caudad 
and dorsad of the left flap of the sinu-auricular valve. 

The conus arteriosus (Plate VI. fig. 18, con.art.) has three longi- 
tudinal rows, each of tliree valves, one row being dorsal, the others 
ventro-lateral. The posterior valves {v^) are pocket-like and very 
thick ; each is connected to tbe posterior face of the corresponding 
middle valve {v") by a strong chorda tendinea, which in the dorsal 
valve takes the form of a vertical membrane attached to the wall of 
the conus along its whole length, while in the ventro-lateral valves 
it is free except at the ends. 

The middle valves (r) are very small and thick, forming knobs 
rather than pouches ; their anterior edges are connected to the walls 
of the conus by several chorda tendinece. The anterior valves (v^) 
are pocket-like and are in close contact with one another at their 
edges, whereas each of the middle and posterior valves is separated 
from its fellow by a considerable interval. The edges of each of 
the anterior valves are produced forwards, forming a firm attach- 
ment, but they have no chordce tendinece. 

There are, as usual, too large coronary arteries placed right and 
left of the conus. 

6. The Vrinogenital Organs. 
The ovary was too much decomposed in specimen C for anything 
to be made of it ; in D it was quite small, so that the specimen 
must have been immature in spite of its size. The oviducts have 
the usual character ; in D there was a considerable dilatation in the 
position of the oviducal gland in one oviduct, but only a very 
shght enlargement in the other. The oviducts open into the exter- 
nal compartment of the cloaca by papilliform terminations (Plate VI. 

fig. 1 7, ovd.apX 

3* 



36 PROF. T.J. PARKER ON CARCHARODON RONDELETII. [Jail. 18, 

The urinary bladder (specimen C) is small and opens into the in- 
ternal compartment of the cloaca by a small papilla (fig. 17, ur.ap.). 

7. The Brain. 

The brain could only be examined in specimen C, in which, as 
already mentioned, decomposition was far advanced before the fish 
came into my possession. As soon as the cranium could be removed, 
a saturated solution of corrosive sublimate was poured into the brain- 
cavity through the foramen magnum, so as to harden the brain in 
situ. Next day the roof of the cranium was sawn off, and the brain 
sketched from above (Plate VII. fig. 20) ; a plate of mica was then 
inserted beneath it, the nerves cut, and the organ transferred to 
strong alcohol with comparatively little shaking. By using these 
precautions, and thanks in great measure to the thickness of the pia 
mater, I was able to make a tolerably accurate examination of tlie 
brain, although all the softer abdominal viscera were hopelessly 
decomposed some days before the brain could be got at. The ence- 
phalon of the fcEtal specimen E was also examined (Plate VIII. 
fig. 29). 

The adult brain is about 13 cm. long from the anterior boundary ot 
the prosencephalon (Plate VII. figs. 20-22, prosen.) to the posterior 
end of the metencephalon ' or medulla oblongata {ineten.) ; to this 
must be added about 12 cm., the length of the olfactory lobes (r/iinen.), 
giving a total length of 25 cm. The greatest width, across the 
cerebrum, is about 3"5 cm. In the foetus (fig. 29) the brain is 
nearly 5 cm. long by 2 cm. wide. 

The main difference between the foetal and the adult brain depends 
upon the elongation, in the latter, of the medulla oblongata and of 
the olfactory lobes. As will be seen by comparison of figs. 20 and 
29, fully one half of the medio-dorsal region of the metencephalon 
is covered by the cerebellum in the foetus, hardly more than one 
sixth of it in the adult. Again, in the foetus, tiie olfactory bnlbs 
are almost sessile upon the prosencephala, their crura being very 
short ; in the adult, on the otlier hand, the rliinencephalic crura 
are of great length. 

In a female specimen of Lanina cornuhica, 135 cm. {4\ ft.) long, 
I find that the brain resembles that of the foetal rather than that of the 
adult Carcharodon ; the olfactory crura are comparatively short, not 
longer than the prosencephala, and fully one half of the metence- 
phalon is covered by the cerebellum. In the fresh brain the optic 
lobes were so completely covered i)y the cerebellum as not to be 
visible in a view from above, but after hardening in corrosive sublimate 
the epencephalon had undergone a slight shrinking, allowing the 
lateral regions of the lobes to be seen. The vertical height of the 
entire brain is great in proportion to its width ; probably in the 
adult Carcharodon the decomposed brain had spread out a good deal 
under the action of gravity. 

In the metencephalon of Carcharodon the restiform bodies (fig. 20, 

' Vide T. J. Parker, " On the Nomenclature of the Brain and its Cavities." 
' Nature,' vol. xxxv. 1886, p. 208. 



1887-] PROF. T. J. PARKER ON CARCHARODON RONDELETII. 37 

rest.) are large and folded. The metacoele or fourth ventricle (mi. 
coe.) is covered by a thick tela vasculosa (tel.vasc.) and its floor is 
marked by five well-marked grooves. 

The epencephalon or cerebellum (e/?en.) is large, divided into lobes 
by transverse sulci, and prolonged forwards so as to cover all the 
median portion of the mesencephalon as well as backwards over the 
anterior sixth of the nietencephalon. It contains a large epiccele 
(cerebellar ventricle), which communicates by a comparatively small 
aperture or ostium with the metacoele. 

The mesencephalon presents dorsally a pair of large optic lobes, or 
optencephala (opten.), and contains a spacious mesocoele (fig. 21, 
(mes.coe.), on the n]iddle cf the floor of which is a small aperture 
(.r) leading into the diaccele or third ventricle (di.coe.). This 
apparently unusual arrangement no doubt indicates that the optic 
lobes have overlapped the posterior portion of the diencephalon. 

The diencephalon {dien.) is short; its cavity, the diaccsle (di.cce.), 
is arched over posteriorly by a narrow bridije of uervous matter, but 
for the most part is covered in only by the thick vascular velum 
interpositum (vel.int.). On its floor is a longitudinal groove (y) 
leading both into the mesocoele and into the iufundibulum, which is 
short and bears a large trilobed hypophysis or pituitary hod j (hi/p.). 
No hsematosac (saecus vasculosus) was ajjparent, and the conarium 
or pineal body was not observed. On the ventral surface of the 
diencephalon are small rounded lobi inferiores (loh.inf.). 

In the foetal brain, as well as in that of Lamna, the diencephalon 
is quite concealed in a view from above, the anterior faces of the 
optic lobes being quite vertical and in close contact with the posterior 
face of the cerebrum (fig. 29). 

The cerebrum is a large, transversely elongated mass, consisting of 
the fused prosencephala or cerebral hemispheres (prosen.), the line 
of junction between which is marked both above and below by a 
distinct groove. Each prosencephalon is also divisible into a large 
dorso-lateral and a smaller ovoidal ventral lobe (fig. 22). 

The cerebrum contains well-developed lateral ventricles or proso- 
coeles (fig. 21, prs.cce.), communicating each by a foramen of Monro 
(for.M.) with a small triangular space, the aula, lying immediately 
cephalad of the diaccele proper, and consisting of the cavity of the 
basi-cerebrnm or unpaired portion of the protencephalon (embryonic 
fore-brain) left by the budding-off of the cerebral hemispheres. 

A choroid plexus {ch.plx.) is continued into each prosoccele from 
the velum interpositum. On the inner wall of the cavity is a large 
ovoidal elevation (a), and a smaller one (b) occurs on its floor. 

The rhineucephalon consists, as already stated, of a greatly 
elongated crus, and of a bulb in apposition with the olfactory sac. 
A cavity, the rhinocoele (rh.coe,) is continued into the crus from the 
corresponding lateral ventricle. 

The first four cerebral nerves present no special features of 
importance. The fifth, seventh, and eighth arise, as usual, close 
together, having between them four principal roots. 

The trigeminal (figs. 20, 22, and 23, v) arises by two roots— an 



38 PROF.T. J. VARKER ON CARCHARODON RONDELETII. [Jail. 18, 

anterior and ventral (fig. 23, v^), former] of two distinct strands, and 
a posterior and dorsaP (v^), which arises mainly from the dorsal 
aspect of the metencephalon (fig. 20), immediately caudad of the 
corpvis restiforme, but also receives a small bundle of fibres arising 
from the lateral region of the metencephalon, in common veith the 
root of the eighth {viii.). The facial (vii.) has one main root formed 
of two strands, the ventralmost of which is intimately united with 
the single root of the auditory nerve (viii.). 

Both dorsal and ventral roots of the fifth divide before leaving the 
skull, so that the nerve passes through the trigeminal foramen in four 
parts (fig. 20), each of which perforates separately the membrane of 
the foramen. 

The sixth nerve (vi.) arises by three distinct roots, the posterior 
of which is very slender and soon unites with the middle root. 

The vagus (x.) is an immense nerve arising by six lateral roots, of 
which the first four and the last two unite to form separate bundles, 
which leave the cranial cavity before joining into a common trunk. 
On the right side (fig. 20) the posterior root is double, and its 
hindmost factor arises at least 1 centim. caudad of the calamus 
scriptorius. 

A short distance cephalad of the origin of the posterior root of the 
vagus there arises from the ventral aspect of the metencephalon a 
distinct though small root (x^), formed by the union of several 
strands. This evidently corresponds with the nerve thus described by 
Balfour ^: — " The main stem of the vagus at a short distance from 
its central end receives a nerve which springs from the ventral side 
of the medulla, on about a level with the most posterior of the true 
roots of the vagus. This small nerve corresponds with the ventral 
or anterior roots of the vagus described by Gegenbaur, Jackson, and 
Clarke (though in the species investigated by the latter authors these 
roots did not join the vagus, but the anterior spinal nerves). Similar 
roots are also mentioned by Stannius, who found two of them in the 
Elasmobranchs dissected by him ; it is possible that a second may 
have been present in Scyllium, but have been overlooked by me, or 
perhaps may have been exceptionally absent in the example 
dissected." 

As the nerve-roots in Carcharodon were made out while the tough 
pia mater was quite intact, I feel satisfied that no other ventral root 
of the vagus was present in my specimen. From the direction taken 
by the nerve it appears to join the vagus, not the spinal nerves ; but 
it was unfortunately severed, as shown in fig. 22, when the brain was 
removed. 

DESCRIPTION OF THE PLATES. 

Plate IV. 
Fig. 1. Cranium of Carcharodon rondelefii, dorsal aspect, X j. 
2. Cranium of Lamna cornubica, dorsal aspect, x 5. 

^ Tliis root propei-ly belongs to the seventh, as shown by Balfour and Marshall. 
' ' Elasmobranch Fishes,' p. 106 (Works, Memorial Edition, vol. i. p. 419). 



1887. J PKOF T. J. PARKER ON CARCHARODON RONDKI.ETIl. 39 

Fig. 3. Cranium of Carcharochn rondektii, ventral aspect, X^. 
4. Cranium of Lamna corimbica, ventral aspect, X |. 

'References fo Figs. 1-4. — aq.fall, aqueductus Fallopii ; font, fontanelle ; 
inf.orh.fl, infraorbital plate ; olf, olfactory capsule ; p.orb.pr, postorbital 
process ; f.oi.'pr, parotic process ; rod"^, dorsal bar of rostrum ; rosP, ventral 
par of rostrum ; vert.ccnt. 1, first_ vertebral body ; i.v., glossopbaryngeal foramen ; 
X, vagus foramen. 

Plate V. 
Fig. 5. Cranium of CarchaTodon rondeletii from the left side, with the upper 
and lower jaws, X5. car.f carotid foramen; k.m, facet for 
hyomandibular ; inf. orb.pl, infraorbital plate ; olf, olfactory capsule; 
op.ped, facet for ojjhthalmic peduncle ; p.orb.pr, postorbital process ; 
rosf^, dorsal, and rost", ventral bar of rostrum ; siip.orb.pl, supra- 
orbital plate ; ii. optic foramen ; Hi. oculomotor foramen ; iv. foramen 
for fourth nerve ; v. trigeminal foramen ; vii. facial foramen. 

6. Anterior extremity of vertebral column of the same showing its 

junction with the cranium, Xj. «', intercalary cartilage ; ra.«, neural 
arch ; vert.ccnt. 3, third vertebral body ; x, vagus foramen. 

7. Part of the neural tube of the same, from the dorsal aspect, X J. /, 

intercalaria ; 11. a, neural arches. 

8. Part of one of the hrenial ridges of tlie same, from the ventral aspect, 

Xj. ife, htemapophyses ; ?, intercalaria ; r, rib. 

9. Hip-girdle and left pelvic fin of the same, X5. a, propterygial 

cartilage; bs.ptg, basipterygium ; dp, cartilage of clasper; p^i, 
pubic bar. 

10. Ventral (so-caUed anal) fin of the same, X |. 

11. Cranium of Lamna cornubica iro\n the left side, with the upper and 

lower jaws, Xj. The reference letters have the same significance 
as in fig. 5. 

Plate VI. 
Careharodon rondeletii. 
Fig. 12. Posterior extremity of vertebral coluiun, X^. H<b, haemapophysis ; 
ha.sp, haemal spine ; pitg, pterygiophore or radial cartilage ; vert, 
cent. 107, 130, & 167, the 107th, 130th, and 167th vertebral bodies. 

13. Vertical section of a trunk vertebra, x4. 

14. Vertical section of a caudal vertebra, X j. 

15. The second dorsal fin, x|- 

16. The stomach and intestine, with the spleen and pancreas, from the 

ventral aspect, X^V- b.d, hile-duct ; card.st, cardiac portion of 
stomach; MJif, - intestine ; pan, pancreas ; ^yAj*;*, pyloric poi-tion of 
stomach ; ret, rectum ; rct.gl, rectal gland ; spl, spleen ; x, sac-like 
dihitations of stomach. 

17. The cloaca with the rectal gland, urinary bladder, and extremities of 

the rectum and left oviduct, X tV- c^\ inner, and cP, outer 
compartment of cloaca ; l.ovd, left oviduct ; ovd.ap, apertm-e of 
oviduct ; ret, rectum ; rct.gl, rectal gland ; ur.ap, urinary aperture ; 
ur.bl, urinary bladder. 

18. The heart from the ventral aspect, the conus arteriosus being 

opened by a longitudinal incision, X:j. aur, aui-icle ; con.art, 
conus arteriosus ; vent, ventricle ; v^, v'-, v^, the three rows of 
aortic valves. 

Plate VII. 

Careharodon rondeletii. 
Fig. 19. Tran.sverse section of the caudal region, Xth- 

20. The brain from the dorsal aspect, the tela vasculosa being removed on 

the left side, nat. size. 

21. Anterior part of the brain with the cavities laid open from above, nat. 

size ; a bristle (.r, y) is passed from the meeoccele into the diaeoele. 



'iO RKV. N. ABRAHAM ON THE [Jail. 18, 

Fig. 22. The brain from the ventral aspect, nat. size. 

23. The roots ol' the 5th, 7lh, and Sth nerves, from the left side, nat. size. 
Eefercnccs to Figs. 20-23. — a, elevation on inner wall of prosoca?le ; aula, 
remains of the cavity of the unpaired cerebral vesicle ; b, elevation on floor of 
prosocoele ; b.opt, basi-opticus ( = ventral portion of mesencephalon); ch.plx, 
choroid plexus; dien, diencephalon ( = thalaniencephalon) ; di.cce, diacoele 
( = third ventricle); f^JC?;, epenceiDhalou (cerebellum); for.M, foramen of 
Monro ; J/i/p, hypophysis cerebri ; mcs.cm, mesoccele ; yneten, metencephalon 
( = medulla oblongata) ; mt.cce, metacoele ( = fourth ventricle) ; opUn, optence- 
phala ( = ojjtic lobes) ; /jtoaym, prosencephala (= cerebral hemispheres), united 
into a single cerebrum ; prs.cce, prosoccele ( = lateral ventricle) ; rhinen, rliinen- 
cephalon ; rJt.cce, rhinocale; tel.vasc, tela vasculosa ; vtl.iut, velum interpositum ; 
i.-x., cerebral nerves. 

Plate VIII. 
Carcharodon rondelctii (foetus). 

Fig. 24. The cranium from the dorsal aspect, nat. size. 

25. The cranium from the ventral aspect, nat. size, font, fontanelle ; 

a.s.c, p.s.c, h.s.c, elevations of the anterior, posterior, and 
horizontal semicircular canals ; h.m, facet for the hyomandibular. 

26. Outer -view of the right auditory capsule, nat. size, h/m, facet for the 

hyomandibular ; h.s.c, elevation for the horizontal semicircular 
canal ; spir.cart, spiracular cartilage. 

27. The ventral region of the branchial skeleton, nat. size. b.hy, basi- 

hyal plate ; b.hr. 2, b.br. 5, basibranchial of the 2nd and Sth arches ; 
h.br. 2, h.br. 4, hypubranchials ; c.hy, ceratohyal ; c.br. 1, c.br. 2, 
c.br. 5, ceratobranchials. 

28. Posterior extremity of the -vertebral column showing the last three 

true centra and the terminal demi -vertebra, X5. 

29. The brain from the dorsal aspect, nat. size. 



3. On the Habits of the Tree Trapdoor Spider of Graham's 
Town\ By the Rev. Nendick Abraham. 

[Eeceived November 15, 1886.] 

Among the very numerous species of Arachnida which are fouud 
through the Caje Colony there are several kinds of Trapdoor 
Spiders. There is a species which, for convenience, I have called the 
Tree Trapdoor Spider, ahout which I wish to give some notes. I 
have been unable to tind any mention of this particular Spider in 
any of my books, or in any I have access to, and it has been until 
now unknown to our local or colonial naturalists, so far as I have 
been able to learn. Thinking it may be known to this Society, 
I have not presumed to name it ; but having very carefully observed 
for many months this wonderful creature, I send you these notes. 

Unlike other Trapdoor Spiders, these build their houses in trees. 
There are certain trees which are more favourable for building- 
purposes thau others, though the trees chosen are various, but in 
each case the trees have a rough bark. The house is a very wonder- 
ful structure, though small, measuring not more thau one and a half 

1 Communicated by Dr. A. Giinther, F.E.S., V.P.Z.S., who stated that the 
Spider in question appeared to be Moggridgia dyeri (O. P. Cambridge, Ann, & 
Mag. Nat. Hist. (4) xri. p. 310, pi. x. 1875). 



1887.] TREE TRAPDOOR SPIDER. 41 

inches in depth. The house is not a burrow, though the spider 
often takes advantage of holes and deep crevices ; but usually it 
is constructed on the surface of the bark, especially if there are 
lumps or prominences near the chosen sjiot. The spider com- 
mences to build by weaving together pieces of bark and other 
substances found in the immediate nei^hbuurhood of the proposed 
house. Tliis part of the work is so skilfully carried out that, when 
complete, it is almost impossible to detect any difference between 
the house and the surrounding bark. I have often placed a piece 
of bark in the hand of a friend and asked that the house might be 
pointed out to me, and this often proves a very difficult task. I 
know of nothing in Nature to surpass this wonderful structure, so far 
as it is an imitation. I have had several of these spiders under 
observation for many months, both in their natural haunts and in 
captivity. Being anxious to know how the doors of their homes 
are constructed, their doors being the most wonderful part of the 
structure, I procured a piece of old stump from a tree and drilled 
several holes into it through the different kinds of surfaces presented 
on the bark. Into each of these holes I introduced a spider ; they 
remainded quite quiet and almost motionless during the day at the 
end of the hole, but on visiting the stump the next morning, I could 
not find the holes until I had made a careful search. I then found 
that a beautiful door had been constructed over each opening, and 
that eac!) door had been made to correspond with the immediately sur- 
rounding surface. One hole had been drilled through a growth of 
lichen ; the door in this instance was made to correspond so perfectly 
that the lichen looked undisturbed, and only after careful inspection 
could the outline of the door be detected. In another instance 
some httle pieces of wood, left by the drill on the border of the hole, 
«ere woven into the door. At first the covering to the opening is 
very thin, like paper, its thickness being increased by numerous lavers 
of silk being added to the inside surface of the door. In this way" the 
sides of the house are strengthened, the whole being very strong 
when completed. In a few trees where circumstances are favourable 
a number of these wonderful houses are to be found, but only by an 
experienced eye. In exploring an old tree some months ago, 1 found, 
high up in the tree, the remains of a large broken branch. This 
branch had been split down, and then torn or cut aveay, leaving a 
trunk attached to the tree, showing a transverse and a longitudinal 
section ; this latter surface of the trunk had been softened by rain 
and atmosphere, and formed a splendid field for these spiders to 
build upon. On a surface measuring 18 inches by 9 I counted 
20 houses, not ail tenanted, some of the spiders having died or met 
with violent deaths at the hands of their enemies. I secured this 
trunk, and now have it in my possession. It is an interesting fact 
that this tree and nearly all the trees on which I have found the 
spiders grow in the High Street of Graham's Town, these trees 
being oaks and " Kaffer-booms." The spiders for years past have 
been able to look out of their little doors upon the busy world, and 
no one knew they were there, until an old friend of mine, who spends 



42 ON THE TREE TRAPDOOR SPIDER. [Jan. 18, 

much time in smoking under one of these trees, saw an open trap, 
and drew my attention to it, and then they could be secreted no 
longer, for I searched every likely tree and made them my special 
care and study for some time. 

I have in my possession two or three houses in which the doors 
have undergone modifications to meet the size of the spiders now 
residing in them. A small individual will sometimes take pos- 
session of the empty house of an adult ; the new comer finds the 
door too large, so constructs another in such a way as to form a 
smaller opening ; thus some houses have two doors : I send you 
a specimen. 

On attempting to lift the doors of these houses, the spiders hold 
them down with great firmness. Knowing that naturalists are un- 
certain as to the means used by the ordinary Trapdoor Spiders for 
holding down the traps, I have taken special care to observe the 
mode adopted by those which inhabit the trees, and I find that the 
hooks of the mandibles, which are barbed, grasp the door, and the legs 
the side of the house. I am quite sure tliat this is the case, for I 
have observed carefully, and in one instance, when the spider held 
on tenaciously, I was enabled to fix open the door and observe with 
a lens, and then to lift out the " fangs," which were buried deep 
in the silken door. I have often found the doors fastened down and 
not held. They are fastened by strong weavings of silk, which must 
be broken before the door can be lifted ; in all such cases the spiders 
do not appear to be active or to assist in keeping down the trap. 
Perhaps at such times the spider is engaged in changing its skin, 
and, in cold weather, hybernating. 

Being anxious to see the spider capture its prey, I put a few grains 
of sugar near one of the doors. Two flies lighted on the sugar, 
and while they were regaling, the trap was thrown open with a slight 
click, the spider darted out, caught one of the flies and retired ; 
the whole transaction was done with such rapidity and dexterity 
that the other fly, though nearly touching the captured one, was 
undisturbed and seemed to be quite unconscious of the fate of its 
companion. I have observed one other capture, and this also was 
carried out with the same extreme rapidity. The spiders are 
probably nocturnal in their habits, though I have never seen them 
out at night, but I know that the work of building goes on during 
the night. The captures I observed were during the day. It may be 
that they work at night to save themselves from detection from some 
of their enemies, and watch for prey both day and night. 

The eggs are placed in a small silken bag at the bottom of the 
nest. When the eggs are hatched, the young live for several months 
a free life in the home of the parent, and are thus protected from 
the ants which infest the trees, until they are strong enough to build 
for themselves ; this they do while they are yet very small, but not 
until they are several months old. The greatest enemies these 
spiders have are the ants ; but the houses are so strong and so much 
like the natural bark that even the ants would not work them much 
damage if they did not catch them, or enter the house accidentally. 
On old trees I have found nearly all the houses without spiders, but 



1887.] DR. SHUFEI.DT ON THE ANATOMY OF CERTAIN AUKS. 4> 

many tenanted by other insects in various stages of transformation. 
Like other spiders, when one meets another there is a fight, which 
often ends in the death of both. 

The spider itself is a very interesting creature. It is about five- 
twelfths of an inch in length ; its legs are short, strong, and flattish. 
The head carries eight simple eyes ; the maxillary palpi of the female 
are leg-like and hooked. There are four stigmata. The colour is 
nearly black. The abdomen is not large in proportion to the rest of 
the body, and bears at its extremity four spinnerets, two large and 
two small. 

I send with this paper specimens of the houses and also of the 
spider. In all cases the houses do not look so well, neither are 
they so perfect as when fresh cut from the trees ; this is partly 
owing to the shrinking and twisting of the bark in drying. If I can 
give any other information respecting this spider, or if it would be 
acceptable to you for me to send other accounts of personal observa- 
tion, I shall be pleased to do what I can, according to the limited 
time I have for this, my favourite pleasure. 



4. Notes on the Visceral Anatomy of certain A.uks. 
By R. W. Shufeldt, C.M.Z.S. &c. 

[Received November 12, 1886.] 

About two years ago the Smithsonian Institution of Washington 
placed in my hands for anatomical description a fine collection of 
bird-skeletons, amounting to nearly a hundred in number, that had 
been collected by American explorers at different times and at several 
localities in the Arctic regions. My researches upon this material 
will quite fill a volume, and are illusLrated by several hundred 
original drawings, the whole being in charge of the Smithsonian 
Institution for pubhcation. When I received this collection it 
was accompanied by a few selected alcoholic specimens of Alba- 
trosses and Auks, sent to me with them in order that I might 
obtain skeletons that were not to be found among the rest of the 
material, my work having chiefly to do with the osteology of the 
groups represented. Among the spirit-specimens of the Auks I 
found one of each of the two interesting forms known to us as 
Brachyrhamphus marmoratus and Syntliliborhamphus antiquus, or 
the Marbled Murrelet and Ancient Murrelet respectively. These 
birds rarely fall into the hands of anatomists in such good condition 
as these were ; and although I only needed their skeletons for the 
purpose I had in view at the time, I nevertheless took the pains to 
carefully remove certain parts of their visceral anatomy, and again 
placing these parts back in the alcohol, I have them now before me 
for examination. 

My surprise was very great to find in these two forms, supposed to 
he very closely related generically, how very different the correspond- 
ing structures and organs occupying the chest and abdomen really 
were. Some of these differences will be readily appreciated by simply 



44 DR. R. W. SHUFELDT ON THE [Jail. 18, 

glancing at the drawings made of them, and which illustrate this 
paper. 

When Forbes was with us and produced his admirable work upon 
the anatomy of the Tubinares which were collected during the voyage 
of H.M.S. " Challenger " ^ he found a great deal that was not only 
unique in the structure of Petrels, but in forms more or less nearly 
related to them. And I am of the opinion that when we come to 
examine carefully into the morphology of Arctic water-fowl, and more 
especially into that of their " soft parts," we shall discover much of 
interest, to say nothing of its importance as throwing light upon tlie 
organization of the types in question, as bearing upon the anatomy 
of the earlier forms of birds ; for it is among these groups, as we 
know, that we find many of the more lowly members of the class in 
])oint of structure and organization. 

This fact was never more forcibly brought to my mind than after 
reading Forbes's investigations and observing the points I am now 
about to describe. 

In S. antiquus (fig. 1, p. 45) 1 find the lower larynx rather broad, 
and somewhat compressed from before backwards. The semirings of 
the bronchial tubes seem to be only partly formed in bone, while the 
last tracheal ring and the pessulus are completely ossified, the latter 
bar being V-shaped on the vertical section, with the apex above. 
"What appears to me as most remarkable about this larynx is the 
mass of fat that overlies it in front, and extends on to its posterior 
aspect, where it becomes thinner. This fat completely covers the 
tracheo-laterales muscles, which are inserted on cither side into the 
middle points of the last tracheal ring. The sterno-tracheales are 
very large and lie embedded in this mass of fat. These are the only 
tracheal muscles present. 

Referring to B. marmoratus, fig. 2, we find the structure of the 
parts to be quite different. In the specimen before me, at least, 
there is an entire absence of fat from this part of the lower larynx. 
The anterior extremities of the lower tracheal ring, which is 
here, too, thoroughly ossified, do not meet so completely as they 
do in S. antiquus, or perhaps, more correctly speaking, this ring is 
roundly notched in front. B. marmoratus has a pessulus of a form 
corresponding very closely to the one described above for S. antiquus, 
but the tracheal tube above it is rather more cyclindrical, and not so 
much compressed from before backwards. The lateral tracheal 
muscles seem to agree quite closely in these two Auks, both as regards 
their size and points of insertion into the mid-lateral parts of the 
last tracheal ring, where they dilate slightly as they become inserted. 
Some considerable difference, however, is to be noted in the sterno- 
tracheal muscles of B. marmoratus, as will be seen in the figure ; they 
are given off much higher up on the trachea iu this Auk, and are far 
slenderer than they are in S. antiquus. 

Unfortunately I neglected to examine the condition of the carotids 

in these two Auks before removing the viscera, as I was intent upon 

not injuring their very brittle skeletons, which had become much 

softened by soaking so long in the partially dissolved fat that encased 

^ W. A. Forbes, Zool. Chall. Exp. vol. iv. pt. xi. p. 1. 



18S7.] 



ANATOMY OF CERTAIN AUKS. 



45 



the bodies of both of them. I will at once observe, however, 
tiiat the form of the heart is quite different in these two birds, being 
not only smaller in S. antiquus than it is in the Marbled Auk or 
Murrelet, but apparently longer, and decidedly more pointed in the 
former than it is in the latter, wherein it is a thicker organ with a 
bluntly rounded apex (fig. 2). 



Fiff. 1. 



Fig. 2. 




Fig. 1. Anterior aspect of the lower larynx, heart, and viscera of SyntMihorham- 
■phiis antifjims. f, fat overlying in front of the lower larynx ; s.t, 
sterno-trachealis muscle of the left side ; h, heart ; I, right lobe of 
liver ; V, left lobe of liver ; g, gizzard. 

Fig. 2. Same parts and aspect of the corresponding organs in Brachyrhamphus 
marinoratus; lettering the same as in fig. 1, with t.I, the left tracheo- 
lateralis muscle. 

The figures are drawn life-size by the author, and are from the specimens of 
the Murrelets lent by the Smithsonian Institution. 

Extraordinary differences are to be observed in the livers of these 
two Murrelets, both as regards form and size. In each the left lube 
is rather the larger, and descends somewhat further into the 
abdomen. But in S. antiquus the hepatic lobes are considerably 



46 ON THE ANATOMY OF CERTAIN AUKS. [Jan. 18, 

longer and narrower than they are in B. marmoratus, and with 
more pointed extremities. In S. nniiquus, too, the connecting band 
of hepatic tissue, joining the two lobes at the back and above, is far 
more extensive than it is in B. marmoratus ; I fail to find any trace 
of a third lobe in either of these Auks. 

Both of these Murrelets possess a large pear-shaped gall-bladder, 
Iving, in either case, beneath the inferior edge of the right lobe of 
the liver. Likewise in each is the spleen well developed ; but this 
organ in S. antiquus is long and subcylindrical in form, while in 
B. marmoratus it is shorter, thicker, and of a decidedly pyriforni 
outline. 

MacgiUivray gives us a very good description, illustrated by three 
figures, of the proventriculus and gizzard of the Little Auk (Mer- 
ffulus alle), winch appears in the eighth volume of Audubon's ' Birds 
of America,' the royal quarto set. In the birds before me I fail to find 
the band of " glandules," arranged as a belt at the extremity of the 
proventriculus, at the entrance of the stomach. Nor is the oesophagus 
so thiu as MacgiUivray found it to be in M. alle : in other particulars, 
however, these Auks seem to be quite similar to it ; for I find the 
inner coat of the elongated proventriculus and the lower part of the 
oesophagus thrown into strong longitudinal rugae or folds, among 
which the surface is thickly studded with minute openings, which 
I take to be the mouths of the glandules. These rugae are con- 
tinuous with similar, longitudinal elevations in the gizzard ; but in 
this latter cavity they are covered by a closely fitting corneous 
structure that readily peals off in the alcoholic specimens, leaving 
the rugae in a condition precisely as we find them in tlie proventri- 
culus and oesophagus. The gizzard and proventriculus are continuous 
and but faintly marked externally by a constriction which shows the 
ending of the latter and commencement of the former, while 
internally, as I say, the definition is made quite sharp by the corneous 
layer of the gizzard. The disposition of the muscles of this latter 
organ are somewhat differently arranged from what MacgiUivray 
gives us in his figure of M. alle. The tendon from which the fibres 
radiated in the Murrelets above described is situated quite laterally, 
and nearly opposite the pyloric exit of the pouch ; while in Macgilli- 
vray's draw ing of the Little Guillemot, already referred to, this gastric 
tendon is centrally located as we see it in Pigeons and other birds. 
Both of my specimens had entirely empty gizzards, the cavities not 
even containing a few grains of coarse gravel, which is not an 
uncommon thing, I believe, in certain Auks. 

The intestines of these Murrelets present us with nothing worthy 
of special remark, and I find a well-developed and large pancreas 
present in each. According to MacgiUivray, in M. alle the rectal 
extremity of the intestinal tube becomes much enlarged and quite 
globular, while a short distance above it we find a pair of caeca of 
no great size. Unfortunately an accident happened to these parts 
in both of my specimens ; but I presume much the same arrangement 
woidd obtain, as, so far as I know, all Auks are thus constructed in 
regard to this part of their economy. 

If hereafter the differences I have pointed out are found to be 



PZ.S.1897.P1,. iZ 




.Sjr!ii.i;lh. 



H3.nl>a/-L iTip. 



I.EUSCARTHMUS APICALIS. 
2.LEPT0POGOT^ OUSTALETI . 



1887.] MR. SCLATER ON NEW SPECIES OF TYRANNIDjE. 4/ 

constant for the lower larynx, the heart, the spleen, and especially 
the livers of these two forms, they certainly constitute very excellent 
generic characters, especially when taken in coiniection with the 
additional ones found to exist in the skeletons. These latter I have 
elsewhere dealt with, but the work at present is in the hands of 
the Smithsonian Institution for publication. 

When good opportunities occur in the future to examine the 
visceral organization of any of the Auks, I trust my fellow-labourers 
will avail themselves of them, and make full comparisons with tlie 
figures I have given above of S. antiquus and B. mnrmoratus, as well 
as test the correctness of my work in the present paper. 

We stand sadly in need of series of alcoholic specimens of Arctic 
water-fowl in the vast majority of our museums. 



5. Characters of new Species of Bii'cls of the Family Tyran- 
nidce. By P. L. Sclater, M.A., Ph.D., F.R.S., 
Secretary to the Society. 

[Eeceived November 30, 1S80.] 

(Plate IX.) 

During the work which I am now engaged upon of preparing the 
Catalogue of the specimens of the birds of the family Tyrannidse in 
the British Museum, I have met with a certain immber of examples 
of this difficult group which are not, so far as I can make out, 
referable to described species. Of these, I beg leave to submit the 
following characters to the Society. 

1. T^NIOPTERA HOLOSPODIA, Sp. UOV. 

Supra ciiierea ; f route et siqjerciiiis. curtis albescentibus ; uropygio 
nigricante ; alls caudague nigricunti-c'mereis ; secundariorum 
externorum apicibus et rectricis utrinque extimce pogonii externi 
margine externa ulbicantibus : suhtus pallide cinerea, in ventre et 
crisso in albidum transiens ; subaluribus albicanti-cinereis ; rostra 
et pedibus nigris : long, tota 8"3, al(£ 6'0, caudce 3'7, tarsi \'7. 
Hub. Bolivia (^Bridges). 
Mus. Brit. 

This species, established on two skins, obtained by T. Bridges in 
Bolivia, in the National Collection, is distinguished from the other 
TcenioptercB by its uniform style of coloration, large size, and long 
wings. The specimens are both in moult. 

2. EusCARTHMus APicALis, sp. nov. (Plate IX. fig. 1.) 

Supra olivaceus ; pileo toto cum capitis laferibus dilute brunneis ; 
laris macula albescente notatis ; alis nigris, harum tectricibus 
dorso concoloribus, primariis et secundariis internis fulvescente, 
secundariis dorso proximis alba latius Umbatis ; cauda cinerea- 
nigricante olivaceo limbata, rectricibus omnibus vitta sub- 
(ipicali nigra, deinde fascia apicali alba prceditis : subtus cine- 
reus ; gula brunnea, pileo concolore ; plaga cervicali magna 



48 MR. SCLATER ON NEW SPECIES OF TYRANNID.E. [Jail. 18, 

antica alba, ventre medio albican te; lateribus olivaceis ; tectri- 
cibus subalaribns albis, campterio fiavicnnte ; rostro brunneo, 
infra dilutiore ; pedibrnpnl/idis: long, tola 3'8, alee I "8, caudcB 
1-6. 
Hah. Brazil. 
Mus. Parisiensi. 

A single skin of tliis well-marked species was presented to the 
Museum d'Histoire Naturelle by M. Peiclioto in 1854. It belongs 
to the group of E. granadensis, and has a similar large cervical spot, 
but is readily distinguishable from all its allies by its pale brownish 
head and the white tips to the tail-feathers. 

I am much indebted to M. A. Milne-Edwards and the authorities 
of the Muse'ura d'Histoire Naturelle of Paris for allowing me to 
bring to England for comparison and identification some of the more 
difficult and obscure examples of the Tyrannidse in that collection. 

3. POGONOTRICCUS GUALAQUIZ^, Sp. nOV. 

" Pogonotriccus gualaquizce, Scl. MS.," Tacz. et Berl. P. Z. S. 
1885, p. 89. 

Similis P. ophthalmico, sed pileo obscure oUvaceo, tectricibus ouri- 
cularibus nan nigro notatis, et colore sttbtus dilutiore diversits : 
long, tola 4'0, ala 1-9, caudce VI . Fem. mari similis. 

Hab, ^quatoria occ. 

Mus. P. L. S. 

The two examples of this species have long remained in my 
collection with the MS. name by which I now characterize them. 
They were obtained by Eraser at Gualaquiza, Ecuador, in January 
1858, and are marked as male and female. Mr. Stolzmann procured 
a single example of the same bird at Mapoto, in the province of 
Ambato. 

4. Leptopogon godmani. 

Supra olivaceo-viridis, pileo obscure cinereo ; loris, superciliis et 
capitis lateribus ulbicante mixtis ; tectricibus auricularibiis flavis 
macula terminali nigra prceditis ; alis nigricantibus flavicante 
bifasciatis, et hoc colore in secundariis externis limb a lis ; ceteris 
remigibus olivaceo marginatis ; cauda cinerascenti-brunnea 
olivaceo limbata ; subtus sulphureo-flavus, pectore et lateribus 
olivaceo tinctis ; tectricibus subalaribns sulphurris ; rostra ob- 
scure corneo, pedibus fuscis : long, tola 4"0, alcB 2'3, caudce '10. 

Hab. ^quatoria orientalis. 

Mus. Salvano-Godmanico. 

Two skins of this species were obtained at Sarayacu, Ecuador, by 
Buckley. It is most like L. superciliaris and L. pcecilotus, but differs 
in its smaller size, yellow wing-bands, and shorter, broader bill. 

5. Leptopogon oustaleti, sp. nov. (Plate IX. fig. 2.) 

Supra olivaceo-viridis, pileo concolore ; linea circumoculari flnva ; 
macula auriculari nigra ; alis caudaque obscure brunneis olivaceo 
limbatis ; campterio jiavo : subtus pallide fiavidus olivaceo 
adumbratus, gula et ventre medio clarioribus ; tectricibus sub- 



188/.] MR. SCLATEll ON NEW SPECIES OF TYllANNID.E. 49 

alaribus pallide sulphureis ; rostro superiore corneo, inferiore 
(libido; j)edibus fuscis : long, tola A 7, alee 25, caudce 2"5. 
Ilah. Colombia interior. 

Obs. Species ciliis oculorum flavis et macula auriculari insigais, 
rostro pauluin latiore quam in speciebus hujus generis typicis. 

I base this well-marked species on a single skin in the Paris 
Museum. There is no label of locality, but the preparation shows 
that it is a " Bogota" skin. 

6. Phyllomyias berlepschi, sp. nov. 

Supra obscure cinerea, dorso olivaceo tincto ; loris albicuntibus ; 
alls caudaque obscure cinereis ; tectricum alurium apicibus et 
secundariorum marginibus externlsangustis albicantibus : subtus 
grisescenti-albida olivaceo Icviter lavata ; gnla et ventre medio 
ferealbis; subalaribus alhis, vixjlnvicante iinctis ; rostro obscure 
fusco ; pedibus nigris : long, iota 4"2, alee 2*2, cauda TS. 

Tlab. Brasilia meridionali-orientalis. 

Mus. P. L. S. 

Obs. Affinis P. griseo-cupillic, sed crassitie minore, et colore 
supra jjallidiore, necnon marginibus tectricum alarium albidis diversa. 

7. Elainea hypospodia, sp. nov. 

Supra fusco-cinerea, cauda concolore ; pilei subcristati macula 
basali alba ; alls nigris, tectricum alurium fasciis transversis 
duabus et secundariorum dorso proximorum marginibus externis 
albis : subtus dilute cinerea, in ventre et crisso et in tectricibus 
subalaribus alba ; rostro fusco ad basin rufescente ; pedibus 
nigris ; long, iota 5'3, alee 2-9, caudcs 2'5. 
Hab. Venezntla. 

In this bird, which belongs to the group of E. payana, there is no 
trace of olive or yellow on the plumage. The single specimen, 
obtained by Goering near Valencia in Venezuela, has been many 
years in my collection under the MS. name now published. 

8. Elainea flavivertex, sp. nov. 

Supra olivaceo-viridis, uropygio dilutiore ; pilei subcristati ma- 
cula basali jiammea ; alis nigricantibus jiavicante bifasciatis, 
necnon secundariis Jiavicante extus marginatis ; cauda fusca 
olivaceo anguste marginata -. subtus cinerea, in gvla albicantior, 
ventre Jiavicante ; subalaribus sulpfiureis ; rostro et pedibus 
obscure corneis: long, lota 4'6, alee 2'3, cauda 2"1. 
Hub. Amazonia superior. 

Obs. Affinis E, gainiardi, sed corpore subtus prsecipue in pectore 
obscuriore, et crista flammea dignoscenda. 

I base this species on a skin obtained by Mr. E. Bartlett on the 
Upper Ucayali, vvhicii has been long in my collection. A similar 
specimen from the same locality is in the British Museum, and a 
third from Elvira (Hauxivell) in the collection of Messrs. Salvin 
and Godman. 

Proc. Zool. Soc— 1887, No. IV. 4 



50 MR. A. Thomson's report on the insect-house. [Feb. 1, 

9. Myiobius subochraceus, sp. iiov. 

Supra sordide olivaceo-viridis, uropygio paulo magis ochraceo ; alls 
nigrlcantibus, tectricum alariiim apicibus latis et secundariorum 
marghiihus externis pallide fiilvis ; cauda obscure cinerea : subttis 
late ochraceus, in rostro medio magis flavesceiis ; subalaribus 
pallide ochraceis ; rostro obscure fusco ; pedibus niyris : long, 
tota A'7, alee 2'. 5, caudcB 2'tJ. 
JIab. Bolivia. 
Mus. S.-G. 

Obs. Affinis M. pulchro, sed crassitie majore et colore subtus 
ochraceo diversus. 

The specimen described is apparently a female of a third species 
of the group of 31. pulcher, distinguished by its large size and the 
uniform ochraceous colouring below. There is just a faint appear- 
ance of a bright colour on the crown, so that the male would 
probably have a concealed orange crest. The bill is rather narrower 
and more elongated than in M. bellus and M. pulcher. 

10. Empidonax ridgwayi, sp. nov. 

Supra obscure olivaceo-viridis, loris et oculorum atnbitu nlbescen- 
tibus ; alis schistaceo-niyris, tectricum ularium apicibus et secun- 
dariorum externorum marginibus externis albescentibus, ochraceo 
vix tinctis ; cauda schistaceo-nigra ; hujus rectricis externce 
poyonio externo albido ; infra sordide albus, in gula media clarior ; 
ventre inferiore et crisso flavicante tinctis; subalaribus albis ; 
rostro superiore obscure corneo, inferiore albido ; pedibus nigris : 
long, tota .50, alee 2" 7, caudce 23. 
JJab. Colombia int. 
Mus. P. L. S. 

Mr. Ridgway marks this bird, which has been submitted to his 
examination, as " probably a new species, most like E. trailli in 
coloration, but \Aith the outer web of the exterior rectrix white, as 
in E. obscurus." I projiose to adopt this suggestion, which I quite 
agree with, and to call the species Enipidotiax ridgwayi, after one 
who has done so much good work in this group of birds. 

The second, third, and fourth primaries are nearly equal and 
longest in this species. The first is shorter than the filth, and yery 
slightly longer than the sixth. 



February 1, 1887. 

Dr. St. George Mivart, F.R.S., Vice-President, in the Chair. 

Mr. F. Day, F.Z.S., exb.ibited a specimen of a hybrid Pilchard, and 
a specimen oi Salmo j)ur])urutus raised in this country. 



A series of specimens of Lepidopterous Insects, which had been 
bred in the Insect-house during the past season, was laid on the 



1887.] MR. A. Thomson's ueport on the insect-house. 



51 



table, and the following report upon the subject, drawn up by 
Mr. A. Thomson, was read : — 

The following species of insects have been exhibited in the Insect- 
house during the past season : — 

Silk-producing Bombr/ces and their Allies. 

Indian. 

Actios sclene. 
Anthertea mylitta. 
Cricula trifenestrata. 



Attacus atlas. 

cynthia. 

ricini. 

jjeruyi. 



Samia cecropia. 
Teleapolyithemus. 
proinetheus. 

Gynanisa maia. 
Anthercea cytherea. 

^ menippe. 

tyrrhea. 



American. 

Actios luna. 
*Dirphia tarquitiia. 



African. 



* Saturnia terpsichore. 

Attacus mythiinna. 
* Actios mimosee. 

Cirinajorda. 



Diurnal Lepidopdera. 
European. 



Papilio podalirius. 

viachaon. 

Thais polyxena. 
Farnassius upollo. 
Euchloe cardamines. 
Vanessa antiopa. 
atalanta. 

* Papilio porthaon. 

* piolice7ies. 

* 



colomia. 



Papilio cresphontes. 



" ajax. 

Smerinthus ocellatus. 

populi. 

Sphiticv ligvstri. 

cvnvolvuli. 

pinastri. 



Vanessa levana. 
^Melitcea maturna. 

Limenitis sibylla. 

Apatura iris. 
* Charaxesjasius. 

Lyccena cor y don. 



African. 



^Papilio nireus. 
'^ deniu/eus. 



morunta. 



American. 



Papilio asterias. 
^ iiirnus. 



Nocturni. 



Arctia caja. 

■ hebe. 

Chelonia villica. 
Lasiocanipa quercifoliu, 
^ pini. 



* Exhibited for the first time. 



52 MR. A. Thomson's report on the insect-house. [Feb. 1, 

Deilephila eiiphorbice. Saturnia pi/ri. 

Chcerocampa porcellus. carpini. 

elpenoi'. *Eurymene dolohraria, 

nerii. Try phcena fimbria. 



Hemaris 7narginalis, Catocala fraxini 

Macroylossa fuciformis. nupta. 

Gallimorpha dondnula. 

Of the Silk-producing jMotlis and their allies, Dirphia tarquinla 
from South America, &\\(\Actias mimosce. Anther cea menippe, Saturnia 
terpsichore, and Attacus mijthimna from South Africa, were all 
exhibited for the first time. Three specimens of D. tarquinia, so 
remarkable for the difference in the size and colour of the sexes, 
emerged in December last ; and I have the honour to exhibit a pair 
this evening. 

The two cocoons of Actias mimoscB, with one cocoon of Attacus 
mythimna, and one pupa each of A. menippe and S. terpsichore were 
brought to England by Mrs. Monteiro from South Africa, where she 
had been collecting insects. The two Actias mimosce emerged in due 
course, but I am sorry to say were cripples. I managed, however, 
in the setting, to get them a little into shape. From the cocoon of 
Attacus mythimna and other two pupa;, fine specimens were obtained. 

During the past season I succeeded for the first time in rearing 
one specimen of the Great Atlas Moth (Attacus atlas) in the Insect- 
house upon ivy. It was quite by accident that I discovered that the 
larvae would eat ivy ; and I was much surprised, in looking over the 
case in which the Atlas INIoths were kept, to find 07ie larva feeding 
upon the leaves of the ivy-plant that was growing at the bottom. I 
then tried the other larvae, which I had feeding upon Berberis 
vulgaris, with ivy-leaves, and found that they ate them freely, and 
seemed to prefer them to the barberry ; but I regret to say that the 
whole of the larvae died in the last stage, although they grew to be 
as large as the one reared. 

The Moth that was reared emerged on the l/th October, 18S6, 
after being in the cocoon about six weeks, and although perfect in 
colour, is one third less in size than those obtained from the 
imported cocoons. 

Attacus pernyi, A. cynthia, and Samia cecropia pair readily in 
confinement; aho Sphinx ligiistri, S. pinastri, Deilephila euphorbice, 
and Chcerocampa elpenor ; and I have reared all from the ova except 
D. euphorbice. 

Of European Diurnal Lepidoptcra, Melitcpa maturna and Ckaraxes 
jasius were exhibited for the first time. The larvae (14) of C. jasius 
were deposited in the Insect-house by Mr. J. H. Leech, F.Z.S., 
previous to his departure for Japan. These larvae were then feeding 
upon Arbutus unedo ; but as the supply of that food failed, 1 tried 
them with Euonymus japonicus, and succeeded in rearing 10 insects 
from the 14 larvae upon it. 

Of African Diurnal Lepidoptera, all the species named were ex- 

* Exhibited for the first time. 



1887.] DR. WINDLE ON THE ANATOiMY OF H\DROMYS. 53 

hibited for the first time ; and all the pupae, with the exception of 
those of Fupilio demohus, were brought home by Mrs. Monteiro. 

Of American Diurnal Lepidoptera, Papilio ajax and Papilio turnus 
were exhibited for the first time. 

Amongst other insects that I obtained last year were a large 
number of the cocoons of, I believe, Thyridopteryx ephemeriformis. 
From these cocoons many male insects emerged and copulated with 
the females, which do not leave the cocoon, and the result was that 
some hundreds of young larva; were produced. Of these only one 
survives, and I exhibit it this evening, in its curious covering. It has 
been reared upon young oak, raised from acorns. When the male 
insects first emerge from the cocoon, their wings are covered with a 
brownish hair, which makes them quite opaque, but on the slightest 
movement of the wings this at once disappears. A full description, 
together with figures, of this insect will be found in the First Annual 
Report of the U.S. States Entomologist, p. H/, by Mr. Charles 
V. Riley. As it is placed amongst the noxious insects by that gentle- 
man, it is perhaps as well, in this case, that I did not succeed in 
rearing more than one of the larva;. 

In conclusion, I take this opportunity of thanking Mr. W. H. 
Edwards, of Coallmrg, West Virginia, through whose kind assistance 
and interest I have been able to obtain many species of Americaa 
insects. 



The following papers were read : — 

1. On the Anatomy of Hydromys chrysogaster. By Ber- 
tram C. A. WiNDLE, M.A., M.D. (Dubl.), Professor of 

Anatomy in the Queen^s College^ Birmingham. (Com- 
municated by Dr. Mivart.) 

[Received December 20, 1886.] 

The following notes are the result of an examination of a specimen 
of the above-named animal, obtained shortly after its death. 

External Appearance. 

Measurements (in centimetres). 

Length from snout to tail Gfi'O 

of tail 29-0 

„ of head 7'0 

Distance from snout to eye 3*3 

J, ,, eye to ear 2*1 

Length of humerus 3*6 

,, forearm 4'2 

„ femur, from apex of great trochanter 5*0 

leg Q-5 

„ hand, to apex of claw of medius 3'5 



54 DR. B. C. A. WINDLE ON THE [Feb. 1, 

Length of vreh of hand, deepest "8 

„ claw of hand, longest '7 

foot 7-8 

web of fdot (deepest at its narrowest part) .. I'.t 

„ claw of foot, longest I'O 

The colour of the back is black with an admixture of golden- 
coloured hairs ; the abdomen is covered with hairs of a dark golden 
colour, a narrow strip of flaxen hairs running longitudinally down 
the body from the lower part of the neck, ceasing at the lower third 
of the abdomen. The tail is black, save for its last fifth, which is 
flaxen-coloured. The hands (vide fig. 3) are armed with moderately 
strono- claws, and the digital interspaces are webbed to a small extent. 
The leet (vide fig. 4) are armed with much stronger claws, and have 
considerably deeper webs in the interspaces. The soles of the feet 
are black, the palms of the hands of a lighter colour, and the dorsal 
surfaces of both are clothed with short golden hairs. The scrotum 
is large and covered with hair ; it contains the testicles, which are 
easily to be returned to the abdomen. 

Muscular System. 
Panniculus. — The dorsal portion extends over the entire back as 
a thin sheet. It is especially strong and well-marked (1) over the 
head, especially the vertex, from which it passes into the checks and 
becomes connected with the roots of the large cheek-hairs; (2) in 
the scapular region and over the latissimus dorsi, with which it has 
some connections ; and (3) over the back of the thighs, where it is 
associated in some degree with the hamstring muscles. The ventral 
part is attached to the inner aspect of the deltoid ridge under the 
deep portion of the pectoralis, and extends downwards from this 
attachment over the thorax and abdomen. 

Muscles of the Head and Necl\ — By the side of that portion of the 
panniculus which passes to the cheek there lies a slender muscle 
which, taking origin from the bone in front of the orbit, ends in a tendon 
which is inserted into the side of the cartilage of the nose (levator alae 
nasi). There is a large elevator of the upper lip, separated from the 
panniculus by the numerous and large branches of the infraorbital 
nerve, and prolonged into the mucous membrane of the roof of the 
mouth as far as the middle line. In front of this is a dilatator naris. 
A small buccinator is present. Levator labii inferioris arises from 
the ujjper surface of the inferior maxilla just posterior to the incisors 
and descends, ex})anding considerably, to the skin below the jaw. 

Masseter consists of three parts : — (1) arises by tendon from a 
prominent tubercle placed at the front of the lowest portion of 
the process of bone extending downwards from the zygoma to the 
superior maxilla and its alveolar portion. The muscular fibres 
connected with this expand and are attached to the edge and internal 
surface of the angle of the jaw. (2) arises from the lower margin of 
the zygoma, and is inserted into the lower jaw from its angle to 
aboutthe position of the roots of the incisors. (3) arises partly from 



1887.] ANATOMY OF HYDROMYS. 55 

the under and inner part of the zygoma witliin the orbit, and partly 
from the superior maxilla anterior to the orbit ; this last portion 
joining the rest by passing between the superior maxilla and the 
zygomatic process just mentioned. Tiiis 3r J part joins the anterior 
part of the 2nd. The remaining facial muscles call for no special 
notice. 

Sterno-mastoid, which is larger than cleido-mastoid, is inserted 
by a small round tendon into tlie mastoid process. Cleido-mastoid 
takes origin from tlie inner jiart of the clavicle, and is attached under 
the last to the mastoid process. 

Steruo-hyoid : both muscles are united into a single sheet without 
trace of median differentiation. The middle part is inserted much 
higher up than the two lateral, viz. in tl>e angle bet.veen the two 
anterior bellies of the digastrics, the lateral fibres being attached 
below the tendinous part of the digastric and near the omo-hyoid. 
The omo-hyoid itself has no central tendon. Digastric has no true 
tendon, the central position connected with the hyoid beino- con- 
stricted and covered on its superficial surface with a few tendinous 
fibres. The two anterior bellies are connected with one another. 

It may perhaps liere best be noted that the two halves of the inferior 
maxilla are extremely movable upon one another, a quantity of 
fibrous tissue intervening at the symphysis so as to form a kind of 
fulcrum by means of which a scissors-like action of the extremely- 
long inferior incisors is obtainable (vide fig. .5). The teeth are 
divaricated from one another partly by the action of the digastrics 
and partly by a transverse iiitermandiiiular nuiscle (fig. .5, a), which 
Hes above the insertion of the digastrics and quite separate from 
them. It is placed at the upper part of the angle between the two 
halves of the maxilla, and is attached to the inferior surface of either. 
Approximation is produced, at least in part, by the masseter and 
especially by the part described above as 1. From this it appears 
that au interval could be produced between the two lower incisors 
during the opening of the mouth which would disappear with its 
closure. Murie and Bartlett \ in a paper on the " Movement of the 
Symphysis of the Lower Jaw in tlie Kangaroo," give an excellent 
account of the mechanism of this movement ni the Macropodidse, and 
quote from Good's 'Book of Nature ' a statement to the effect'that 
a similar movement takes place in Mus mariiimus, the African rat. 
In Hydrcmys there is no such development of the transverse fibres 
of the orbicularis oris as the above authors describe in Halmaturus 
bennettii ; whilst the intermandibular muscle above mentioned is 
quite distinct from any of the other inframaxilLiry muscles, all of 
which are present in addition. The amount of divergence possible 
would be from g to j inch. 

Muscles of Shoulder-girdle and Upper Extremity. — The two 
portions of the trapezius are quite distinct, the origin of the 
lower being partly from the lumbar fascia. There is a large 
occipital rhomboid, and rhomboidei major and minor form a sin»le 
sheet without sej.aration. Orao-cervicalis arises from the transverse 

1 P. Z. S. 1806, p. 28. 



56 DR. B. C. A. WINDLE ON THE [Feb. 1, 

process of the atlas, and is inserted into the spine of the scapula and 
acromion process as far as its apex. Pectoralis major consists of 
two layers folded in upon one anotlier so as to present a rounded 
anterior border without any trace of division. The superficial portion 
arises from the clavicle and from the sternum as low down as the 
upper part of the xiphoid cartilage. This part is inserted (1) as 
usual, (2) into a common tendon with the deltoid. Tbe deeper 
portion arises from the cartilages of tlie true ribs from the third to 
the last. This is inserted by two distinct slips, one of which passes 
to the tip of the acromion process, and the second to the fascia of 
the shoulder-joint and to the humerus external to the biceps, and as 
low down as to the upper edge of the attachment of the superficial 
portion. As has been above observed, there is no trace of the du- 
plicity of these muscular sheets at the edge ; in fact it is only by 
dissecting carefully through the outer that the inner is reached. 
This inner sheet is obviovisly the pectoralis minor, and the condition 
present is one of extreme fusion of the two pectoral muscles ; or, per- 
haps better, of complete tucking-in of the p. major to form p. minor. 

Subclavius is strong and well-marked, a fact which corresponds 
•with the comparatively small and freely movable clavicle. Sarratus 
magnus and levator anguli scapulas form, a single undivided sheet. 
Latissimms dorsi sends down a fairly broad but Tery thin latissimo- 
condyloideus to tlie olecranon. Coraco-brachialis is inserted into 
the humerus (1) in the usual position ; (2) from this point as far 
down as the upper part of the internal condyle. The long head of 
the triceps is verv large and arises from rather more thair one third 
of the axillary border of the scapula. Flexor profundus digitorum 
consists of two parts which unite under the annular ligament : the 
first arises from the internal condyle, the second from the radius, ulna, 
and interosseous ligament. There are three lumbricales, passing to 
minimus, annularis, and medius. Extensor communis digitorum 
sends a slip to each digit. Extensor indicis supphes that digit alone ; 
and extensor minimi digiti sends tendons to minimus and annularis. 
Pollex has one extensor. There is no supinator longns. 

Minimus has an abductor arising entirely from the pisiform, an 
opponens and a flexor brevis. The last arises from a small ossicle 
imbedded in the palmar fascia, slightly to the radial side of the 
centre of the ]>alm and at its proximal portion. From this also 
arise the few fibres representing flexor brevis pollicis. This last 
diminutive digit has also on its outer side a few fibres representing 
abductor and opponens, and on its inner side an excessively rudi- 
mentary adductor. Minimus has an interosseous on its radial side ; 
and each of the remaining digits has a pair lying in the same plane 
on its palmar surface. 

Abdomi7ial Muscles. — At the upper jtart of the abdomen, the 
three usual lateral muscles are present and distinct ; at the lower 
portion, as the fibres of the internal oblique and transversalis run 
parallel and are closely connected w ith one another, there can hardly 
be said to be any true differentiation between them. From the 
aponeurosis of the external obHque a sheet of fascia passes down upon 
the large funicular process of peritoneum containing the testicle. 



I8»7.] ANATOMY OF HYDROMYS. 5/ 

This represents the intercoluninar fascia of human anatoni}'. From 
the lower border of the combined internal oblique and transversalis 
a number of muscular fibres pass on to the same process and sur- 
round it as a series of rings nearlj^ to its lower end. There is a large 
erector penis overlying the crus of either side, but no transversus 
perinei. The bulbus urethrre is double, and each half is overlaid bv 
a muscular sheet which joins its fellow of the opposite side in a 
raphe on tlie lower surface of the penis ; posteriorly the two halves 
unite behind the rectum, round which tube they form a kind of 
sling. In the angle betw^een this last muscle (accelerator urinjp.) 
and erector penis lies Cowper's gland. The psoas and iliacus 
muscles are both large and differ in no respect from the normal. 
There is no psoas parvus. Rectus abdominis is attached to the 
second rib. 

Muscles of the Lower Extremity. — The exterior of the buttock is 
covered by a large sheet of muscle arising from (!) crest of ilium, 
(2) under the anterior superior spine of the ilium, (3) by means of 
an aponeurosis from all the vertebras from the last lumbar to the 
last sacral inclusive. It is inserted into (1) the third trochanter at 
the middle of the femur; (2) the larger part into the fascia on the 
outer side of the thigh and leg and the upper part of the patella. 
As there is no separate tensor vaginse femoris or sartorius, this 
muscular sheet would appear to represent these two in fusion with 
gluteus maximus. Gluteus medius is very large ; its anterior fibres 
are inserted into the outer edge of the great trochanter, its posterior 
into the femur inferior and anterior to this process. Gluteus mini- 
mus arises from the concavity of the ilium, and is inserted by tendon 
into the upper part of the great trochanter. There is no separate or 
intrapelvic ])yriformis; part of the fibres of the gluteus minimus 
arising from the edge of the sacrum close to the sacro-sciatic foramen 
aj)pear to represent this muscle. 

Biceps is very large, and arises (1) superficially from the up|)er 
caudal vertebrae by fascia ; (2) deeper, from the' tuberosity of the 
ischium. The two parts unite, and are inserted (1) by fascia into 
the outer part of the patella; (2) by tendon into the process near 
the head of the fibula ; (3) by fascia into the whole of the front of 
the leg as far down as the back of the heel. Thus the entire of the 
thigh and leg below gluteus maximus is covered by this large mus- 
cular sheet. 

Semitendinosus is single-headed and arises from the tuberosity and 
adjacent portion of tlie ischium ; it is inserted into the crest of the 
tibia below the gracilis. Semimembranosus, which is very much 
smaller than either of the other hamstring muscles, is inserted 
into the upper part of the posterior aspect of the internal condyle of 
the femur. 

Rectus femoris has a single tendon with a double attachment, viz. 
under the acetabulum and to the margin of the ilium. There is a 
scansorius arising from the entire of the anterior edge of the ileum. 

Pectineus consists of two distinct parts — (1) Internal, which is thin 
and arises from the inner part of the linea ileo-pectinea, some of 
its fibres underlying the outermost of gracilis ; this portion is inserted 



53 DR. B. C. A. WINDLE ON THE [Feb. I, 

by a flat tendon into the back of the femur at the junction of its 
lower aiid middle thirds. (2) External, which is much thicker and 
rounded, arises from a prominent tubercle near the centre of the linea 
ileo-pectinea, and is inserted into the femur from the lower part of 
the lesser trochanter to the upper border of the internal part of the 
same muscle. 

Gracilis is very large, and arises from the inner part of the linea 
ileo-pectinea, from the pubic crest, symphysis pul)is and ramus of that 
bone, covering in the other adductors ; it is inserted into the crest of 
the tibia, occupying nearly its up[)er iialf. Adductors longus, magnus, 
brevis, and quadratus are present ; the lower fibres of niagnns pass 
down as low as the head of the tibia. 

Gastrocnemius is large and its heads are without sesamoids. It 
is joined by soleus, which is small, and fibular only in its origin. 
There is no separate plantaris, but the posterior part of the tendo 
Achillis passes over the back of the os calcis, to which by far the 
greater part of the same tendon is attached, to the sole of the foot. 
With this tendon are connected on its superficial surface a number 
of muscular fibres, from which and from a slight continuation on the 
deep surface of the fibres of the plantar portion of the tendo Achillis 
arise the four perforated tendons of the toes. There is no flexor 
longus hallucis as a sef)arate structure. A large muscle occupies the 
whole of the internal and posterior portion of the leg : this ends in a 
single large tendon, which divides into five slips for the toes. Tibialis 
posticus is a small muscle with a long tendon ; its belly lies under the 
upper part of flexor longus digitorum. There are peronei longus, 
brevis, quinti, and quarti. Tibialis anticus and extensor longus 
digitorum, which last has four terminal tendons for the four outer 
toes, are much fused. There is a small bnt distinct extensor 
proprius hallucis. Extensor brevis digitorum sends tendons to the 
four outer toes. There is a large popliteus. Hallux has an adductor 
which arises from the middle of the inferior surface of the os calcis, 
and ends in a long tendon which is inserted into the inner side of the 
head of the metatarsal ; an opponens and a strong flexor brevis. 
Minimus has a strong abductor and an interosseal flexor brevis. 
Each of the remaining digits has a single muscular mass lying on the 
plantar aspect of its metatarsal without other trace of division 
than an incomplete median longitudinal fibrous intersection. This 
muscle is in each case inserted into the base of the first phalanx and 
into the superior surface of the fibro-cartilage lying over the meta- 
tarso-phalangeal articulation. 

Nerve Plexvses. 
JBrachial Plexus (fig. 1). — The fourth cervical nerve divides into 
two portions, from the upper and smaller of which is derived the 
greater part of the phrenic. The lower portion joins the 5th, which 
has previously given a filament to the phrenic. The combined cord 
formed from 4 and 5 breaks up into four branches — supra- and sub- 
scapular (smallest), circumflex and musculo-cutaneous. Circumflex 
gives off" a branch of communication to the upper part of the tith 
uerve and also two subscapular branches. Musculo-cutaneous gives 



1S87.J 



ANATOMY OF HYDROMYS. 



59 



off the greater portion of the anterior thoracic nerve. The Gth nerve 
divides into two portions, an npper and lower. The combined cord 
formed by the 7th cervical and 1st dorsal also divides into two 

Fig. 1. 







M.Sr 



Cervical and brachial neryes. (Diagrammatic.) 

1-7. Cervical nerves; LB. 1st dorsal nerve; Sp.Sc. Suprascapular ; Sb.Sc. Sub- 
scapular ; CfLr. Circumflex; Tl/.C. Musculo-cutaneous ; Tr. Nerve to tri- 
ceps; M.Sp. Musculo-spiral ; U. Ulnar; M. Median; C'u. Internal cuta- 
neous ; Phr. Phrenic ; Tk. Anterior thoracic ; M.M. Muscular branches. 

portions, an anterior and a posterior. The upper part of the sixth, 
having received the branch from the circumflex, joins the anterior 
part of 7 + 1, and the combined trunk becomes musculo-spiral, having 
previously given oflF branches to the triceps and other muscles. The 
lower division of 6 joins the posterior of 7+ 1, having first given off 
a filament which joins the anterior thoracic. The large trunk thus 
formed, having given off some muscular branches, splits up into ulnar, 
median, and internal cutaneous. 

Lumbosacral Nerves (fig. 2).— The 1st lumbar forms the ilio- 
hypogastric and the 2nd the ilio-inguinal. These nerves are 
connected with one another by a communicating branch, but have no 
junction with those below. The 3rd gives off genito-crural, a 
branch of communication with 4, and in conjunction with a branch 
from this last forms external cutaneous. Tlie remaining portion of 
4 with a branch from 5 forms a common trunk which divides into 
obturator and anterior crural. The great sciatic is formed by the 



60 



DR. B. C. A. WINDLE ON THE 



[Feb. 1, 



greater part of the 5th lumbar, the whole of the 1st sacral, and a 
twig from the 2ncl. The pudic nerve is formed from the remaiuiug 
portion of the 2nd. 

Fix. 2. 




E.c 



Lumbar and sacral nerves. (Diagrammatic.) 

I-hy. nio-liypogasfric ; II')'g. Ilio-ingiiinal ; E.C. External cutaneous; Ac, 
Anterior crural ; Ob. Obturator; 6'c. Genito-crural ; G.Sc. Great scaitic ; 
P. Pudic. 

Thorax. 

The chief points of interest are as follows: — (1) Triangularis 
sterni is very large and well-marked. It ascends nearly as high 
as the upper margin of the sternum. (2) The diaphragm is in 
most respects in no way different from the ordinary condition. Its 
central tendon is, however, triradiate, consisting of three strips of 
fibrous tissue arranged somewhat like an arrow-head ; one of these 
is directed towards the sternum, a second backwards and to the right, 
a third backwards and to the left, and the interval between these 
last is muscular. 

(3) Lungs. — The left has three lobes, divided in a way similar to 
that of the right lung in the human subject. The right has five 



1887.] 



ANATOMY OF HYDROMYS. 



61 



lobes, the supernumerary pair being placed behind and below the 
others. 

Alimentary Canal and Appendages. 

Teeth. — The dentition is as described by Owen', I. -,, M. \, the 
total number thus being smaller than that of any other Rodent, 
The incisors closely resemble tliose of the Common Rat. 

The upper incisors are stouter and shorter than the lower ; they 
are set closely together, their edges meeting at their inner angles, so as 

Fi-. 4. 





Fis; 5. 




Fig. 3. Left band of Hydromys chrysogaster. (Natural size.) 
Fig. 4. Left foot of Hydromys chrysogaster. (Natural size.) 
Fig. 5. Upper and lower incisoi-s of Hydromys chrysogaster: a, iuter- 
maudibular muscle ; b, tongue. 

to enclose a wide angle {vide fig. 5). The portion projecting beyond 
the alveolus is 1-05 cm. in length. The first upper molar is a 
large tooth, consisting of two segments of nearly equal size and a third 
of lesser magnitude {vide fig. 6, b and c). It possesses three large 
fangs, one at either extremity and the third at the middle and on the 
lingual edge. At the opposite side of the tooth to this last are three 



' Odontography,' vol. i. p. 410, and Comp. Auat. of Vert. vol. iii. p. 300. 



62 



DR. B. C. A. WINDLE ON THE 



[Feb. 1, 



excessively small points, each of which fits into a corresponding 
depression in the alveolus {vide fig. 6, c). 

The second upper molar looks like one of the divisions of the 
first with an additional cusp attached to its antero-iiiternal corner. 
It has three roots, two posterior with their surfaces j)laced external and 
internal, and one anterior with its surfaces anterior and posterior. The 

Fig. 6. 





Eydromys cliry&r,c;aster: «, lower jaw; t, iipj^er jaw ; c, upper molar (enlarged). 

Fig. 7. 




Stomach oi Hydromys chrysogaster : oe, cardiac orifice; py, pyloric orifice. 

lower incisors {vide figs. 5, 6, a) are of greater length (r55 cm.) and 
of about the same breadth, but of less thickness (0"25cm. as compared 
with 0"4 cm.) than the upper. Their power of approximation, due to 
the mobility of the two halves of the inferior maxilla, has already been 



18S7.] 



ANATOMY OF HYDROMYS. 



63 



dwelt upon. The lower first molars consist of two nearly similar 
segments f)laced one in front of the other ; and the second are much 
the same in appearance but smaller (vide fig. 6, a). 

Tongue. — 4-() cm. in length ; it has an average breadth of TO cm. 
It possesses a single large circumvallate papilla, situated at the centre 
near the posterinr part. Tiiere is a small group of folia ou either 
side in front of the anterior pillar of the palate. 

Intestinal Canal. — The stomach is simple in structure, and the chief 
points to be remarked upon are the increase in size of the pyloric 
portion and the decrease in length of the part corresponding to that 
named in the human subject the lesser curvature (vide fig. 7). The 
small intestines measure 2 m. 89-5 cm. ; the large 27-8 cm. ; and the 
caecum about 7*0 cm. This latter portion is quite simple and only 
slightly cnrved upon itself {vide fig. 8). 

Fig. 8. 




Caecum of Hydromt/s chrysog aster. (Natural size.) 
Co. Colon ; n. Ileum ; Coc. Csecura. 

Liver. — This organ possesses six lobes. The right lateral is short 
and permits the caudate, which is comparatively long, to be seen 
U|)on the upper surface. It appears between the right central and 
lateral lobes. There is no gall-bladder {vide figs. 9 and 10). 

Genitalia. — The vesicnlae seminales are very long and narrow, 
extending a considerable distance above the bladder. Their length 
is 5'6 cm., average breadth about 1"0 cm. The testes are large and 
■were, when the animal was examined first, placed in the scrotum. 
The length of each is 4'2 om., circumference 62. In life these 
measurements would probably be larger, as they were made after the 
animal had been in spirit and water for a day or two. The globus 
major and minor are very distinct and united by a single tube which 
is perfectly straight. The length of major is AS cm., minor 2'1. 
There is a small os penis (I'l cm. in length), consisting of a some- 
what square-shaped base with a tapering anterior portion. From 
each side of the base a small process descends, each of which lies on 



64 



ON THE ANATOMY OF HYDROMYS. 

Fis. 9. 



[Feb. 1, 




Superior surface of liver of Hyclromys chri/sogasier. 

L.L. &.L.C. Left lateral and central lobes; E.L. & E.G. Eight lateral and 
central lobes ; G. Lobulus caudatus ; <S'. Lobulus Spigelii. 




Tinder surface of liver. 

L.L.kL.C. Left lateral and central lobes; E.L. & E.G. Eiflit lateral and 

central lobes ; C. Lobulus caudatus ; S. Lobulus Spigehi, 



P.Z.S.1887.P1.X. 















W.Purkiss itK . 



rlsnaarL nnp , 



PHYTOPHAGOUS COLEOPTERA 3E CETLGN 



P,Z.S.1887, P].XI 












Y.rurJsss iu,n 





Hsiikart imp 



PHYTOPHAGOUS COLEOPTEM OF CEYLON 



1887.J ON THE PHYTOPHAGOUS COLEOPTERA OF CEYLON. 65 

one side of the urethra; at the anterior extremity of the hone is 
placed a small rod of cartilage S cm. in length, which gives in the 
undissected state the impression that the l)one is jointed in the 
centre. 

There are none of the appendages connected with the glans wliich 
are present in some of the Rodentia. Cowper's glands are each of 
about the size of half a pea ; their ducts are of considerable length. 

I wish to express my acknowledgments to Mr. Ilodson for his 
kindness in executing the accompanying drawings of the teeth of 
this Rodent. 



2. Descriptions of the Phytophagous Coleoptera of Ceylon, 
obtained by Mr. George Lewis during the years 1881- 
1882. By Martin Jacoby, F.E.S. 

[Received December 21, 1886.] 
(Plates X. & XI.) 

Since the year 1866, when Motschulsky published his descrip- 
tions of Ceylonese Coleoptera in the 'Bulletin de Moscou,' which 
included a good many species of the family Phytophaga, only solitary 
new species of that family have been made known from Ceylon. 
The present collection, obtained by Mr. George Lewis during the 
years 1881-82, although not very large in regard to numbers, is 
nevertheless remarkable and interesting on account of the many 
new genera which it contains, belonging principally to the sub- 
families HalticincB and GalerucincE, in which the immense numbers 
of forms which are nearly always found in every fresh collection of 
importance, and which cannot be placed in any of the already known 
numerous genera, add not a little to the difficulties experienced by 
the monographer in their determination. Motschulsky's descriptions 
are unfortunately in many instances unrecognizable, being applicable 
to closely allied species and too short ; and it is therefore probable 
that several of the species here described as new are identical with 
one or the other of Motschulsky's species ; but even in that case 
their redescription will assist in their better recognition. 

If one may judge by the present collection, the Coleopterous 
fauna of Ceylon seems yet to offer a rich field to a careful explorer, 
in interesting and beautiful forms. 

Lema ceylonensis, sp. nov. 

Fulvous ; antennae black, the two basal and the two apical joints 
fulvous ; elytra metallic green or blue, with a deep fovea below the 
base, finely punctate-striate. 

Length 2 lines. 

Head entirely fulvous and impunctate, the space above the eyes 
moderately swollen and divided by a shallow groove. Antennae 
rather more than half the length of the body, the third and fourth 

Proc. Zool. Soc— 1887, No. V. 5 



66 MR. M. JACOBY ON THE [Feb. 1, 

joints of equal length, the following move elongate. Thorax sub- 
quadrate, not longer than broad, moderately constricted at the sides, 
the basilar sulcation deep ; surface with a few scarcely visible punc- 
tures. Scutellum fulvous. Elytra of a bright metallic yellowish 
green or blue, with a very deep fovea below the base, near the 
sutural margin, finely and rather remotely punctate-striate, the 
punctures obsolete towards the apex. The legs and tarsi fulvous, the 
posterior tibise slightly curved. 

Bogawantalawa, 4900-.5200 feet. 

It will not be very difficult to recognize this species amongst the 
small metallic blue forms of Lacordaire's first section, with uninter- 
rupted ninth elytial stria; the colour of the antennae, which have 
sometimes the first three joints and the last fulvous, or the underside 
of the three or four terminal joints of tliat colour, the deep elytral 
fovea, and the fine and remotely placed elytral punctures, will help 
to distinguish L. ceylonensis ; the lateral margin is accompanied by 
a deejier row of punctures and is costate towards the apex, but the 
other interstices between the punctures are perfectly flat, the reverse 
being the case with most of the allied species. 

Lema fulvicornis, sp. nov. 

Subquadrate-ovate ; fulvous ; labrum black ; thorax impunctate ; 
elytra dark blue, deeply and regularly punctate-striate. 

Length 3 lines. 

Head with the interocular space strongly swollen and finely punc- 
tured, the lateral grooves very deep ; the labrum and upper part of 
the clypeus black, the former with some transversely placed punc- 
tures. Antennae rather more than half the length of the body, 
entirely fulvous, the fourth joint very slightly longer than the third, 
the following joints elongate, cylindrical, and not increasing iu 
thickness. Thorax scarcely longer than broad, deeply constricted at 
the sides, the basilar groove also deep, the anterior angles slightly 
pointed but not tuberculate ; the surface convex and swollen, without 
any punctures. Elytra broad, subquadrate, the shoulders moderately 
prominent, the base scarcely depressed, the punctuation deep and 
not very closely placed anteriorly, much more clijse and diminishing 
posteriorly, where the punctures themselves are placed in striae and 
tiie interstices longitudinally costate. Underside fulvous, clothed 
with yellow pubescence ; legs robust, entirely fulvous. 

A single specimen. 

L. fulvicornis seems closely allied to L. prceclai-a, Clark, but 
differs in its much smaller general size, in the colour of the head 
and of the elytra. 7>. ci/aniiiennis, Lac, is larger and the interstices 
between the punctures of the elytra are finely punctate. The 
present species may be further known by its broadly subquadrate 
shape. 

Lema crassicollis, sp. nov. 

Blackish blue below; upper part of head, the thorax, and the 
last two joints of the antennae fulvous; elytra metaUic blue, deeply 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 67 

foveolate below the base, fiaely punctate-striate, a small spot above 
the shoulders fulvous. 

Var. The lateral margia and the apices of the elytra fulvous ; 
femora testaceous below. 

Length 2 lines. 

Head finely punctured at the vertex, the parts of the mouth 
black. Antennae more than half the length of the body, black, the 
last two joints fulvous, third and fourth short, of equal length, the 
fifth joint double the length of the preceding. Thorax not longer 
than broad, the anterior portion strongly swollen, its angles tuber- 
culiform when seen from above, each angle furnished with a single 
hair ; basal groove very deep ; the surface entirely impunctate. Elytra 
with a deep fovea below the base near the suture, the basal portion 
above it raised ; the surface rather deeply punctate-striate, the punc- 
tures not very closely placed and diminishing towards the apex, the 
interstices slightly transversely wrinkled, longitudinally costate near 
the r.pices ; just above the shoulders at the basal margin, a small 
fulvous spot is placed. Legs black, the underside of all the femora 
fulvous. In the variety the entire lateral and apical margin of the 
elytra is of that colour. 

The elytra iu this species have the same deep fovea as in L. cei/- 
lonensis, but their punctuation is much stronger and the antennae 
have no fulvous basal joints ; the thorax also is much more swollen 
anteriorly ; and the general coloration is ditferent. 

Lema difficilis, sp. nov. 

Below bluish black ; head and thorax fulvous ; antennae black ; 
elytra metallic blue, obsoletely depressed below the base, regularly 
and strongly punctate-striate, the interstices costate near the apices ; 
legs fulvous, stained with piceous. 

Var. Elytra fulvous, a sutural and lateral longitudinal band blue. 

Length 2 lines. 

Head with some fine punctures when seen under a strong glass ; 
the vertex but little swollen, with the usual central groove, this 
latter short and superficial. Antennae more than half the length of 
the body, black, the two lower joints stained with fulvous below, 
third and fourth of equal length. Thorax not longer than broad, 
the anterior angles pointed wlien viewed from above, the sides 
rather deeply constricted ; surface with a deep basal transverse 
groove ; the disk with two longitudinal rows of more or less distinct 
punctures, the sides with some punctures anteriorly only. Scutellum 
fulvous. Elytra with the base slightly raised and depressed below, 
the punctuation strong and regular, but more deeply impressed at 
the basal portion. Legs piceous, more or less stained with fulvous. 

Dikoya. 

L. difficilis cannot be considered a small variety of L. coroman- 
deliana, on account of the absence of the anterior thoracic groove. 
L.javana, Lac, is distinguished by the blue head according to the 
author's description ; in the present insect it is entirely fulvous ; the 
thorax also is almost broader than long, and the anterior angles are 

5* 



68 MR. M. JACOBY ON THE [Feb. I, 

projected into a small tooth or tubercle vvlien viewed from above. 
In the variety the elytra are blue, with a broad discoidal and a 
narrow lateral longitudinal baud, which iu another specimen are just 
indicated, thus proving the identity with the type, with which they 
agree iu every other respect : this variety seems to be closely allied 
to L. 1-iifo-ornata, Clark, in which the elytra, however, are black 
and the underside fulvous. 

Crioceris semipunctata, Fabr. 

Kitulgalle, 1 700 feet. 

Two specimens, evidently females, were obtained by Mr. Lewis, 
which agree very nearly with the Malayan forms ; the antennse are 
very short and robust, and the terminal joints are transversely quad- 
rate ; the punctuation of the elytra is only visible anteriorly, no 
traces of any punctures are seen below the middle ; below the 
shoulders, close to the lateral margin, a short and very deep row of 
punctures, interrupted in the middle, are seen — a character which I 
do not find mentioned by Lacordaire, with whose description the 
Ceylonese specimens agree in all other respects. 

DiAPROMORPHA QUADRIPUNCTATA, Sp. UOV. 

Pale fulvous, finely pubescent below ; the antennae black, the 
four basal joints fulvous ; thorax very minutely, elytra closely and 
distinctly punctured, each elytron with four black spots (2 2). 

Length 3 lines. 

Head impunctate, flattened between the eyes, and with an obsolete 
triangular depression ; eyes slightly notched at their inner margin. 
Antennse short, the third joint shorter and much thinner than the 
second, the fifth and following joints transverse. Thorax three 
times as broad as long, the sides slightly rounded and narrowed in 
front, the posterior margin straight at the sides, broadly truncate 
at the middle ; surface with an obsolete transverse groove at each 
side, very finely and rather distantly punctured. Scutellnm impunc- 
tate, its apex slightly raised. Elytra subcylindrical, distinctly lobed 
at the sides near the base, very closely and distinctly punctured, 
with some obsolete longitudinal smooth lines ; each elytron with two 
black spots, placed transversely below the base and two others below 
the middle, in a line with the anterior spots, the outer one, however, 
being placed slightly higher ; the first joint of the posterior tarsi 
distinctly longer than the second. 

There is only a single specimen, evidently a female, before me. 

Chlamys pallifrons, sp. nov. 

Brownish black ; basal joint of the antennse testaceous ; lower 
part of the face flavcus ; thorax elevated behind, closely granulate- 
punctate, spotted with fulvous in front and at the base; elytra dis- 
tinctly punctured, with an oblique ridge at the middle of the disk, 
a shorter one at the shoulder, and several tubercles at the sides and 
apices. 

Length 1 line. 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 09 

Head closely granulate-punctate, black, the lower part of the 
face flavous. Antennae dentate from the sixth joint, the two or 
three lower joints flavous, the rest black. Thorax strongly raised 
posteriorly into an undivided hump, extremely closely and finely 
granulate-punctate, black ; the hinder portion of the elevation as 
well as some indistinct spots placed anteriorly, fulvous ; a more 
distinct similarly coloured spot is placed at each side of the eleva- 
tion. Scutellum transverse, its posterior margin emarginate. 
Elytra slightly constricted at the middle, of a more brownish colour, 
more strongly punctured than the thorax ; each elytron with an 
indistinct ridge from the middle of the base to the apex and joined 
by a shorter one commencing at the shoulder; the subsutural ridge 
is joined to the suture at the middle by a short transverse elevation, 
and ends near the apex in a strongly raised longitudinal tubercle ; 
three other small and rather indistinct tubercles are seen within the 
elytral constriction at the sides ; the base of the femora is obscure 
fulvous ; the rest and the underside blackish ; the suture is dentate 
through its entire length. 

A single specimen. 

C. palUfrons resembles in general structure C. spilota, Balv, but is 
only half the size ; the thorax is more finely punctured and devoid 
of raised lines and tubercles, while those of the elytra are also much 
less distinctly raised and the interstices much more finely punctured. 

EXEMA CEYLONENSIS, Sp. UOV. 

Black ; head strigose ; thorax strongly raised behind, with six 
elevated longitudinal ridges ; elytra strongly punctured, each elytron 
with an elevated ridge and about nine or ten tubercles ; pygidium 
with three longitudinal carinations. 

Length 1 line. 

Head closely covered with longitudinal strigse ; antennse black, 
the sixth and following joints transverse. Thorax strongly raised 
into a pointed hump, rather closely and distinctly punctured ; each 
side with three longitudinal ridges, the middle pair running nearly 
parallel and not extending quite to the anterior margin, the second 
ridge strongly sinuate, the third the shortest and only extending to 
the middle of the thorax ; there are a few short and obsolete ele- 
vations placed between the above-named ridges, connecting the 
latter here and there with each other. Elytra more deeply punc- 
tured than the thorax ; each elytron with the following tubercles : 
one at the middle of the base, one at the shoulder, and another near 
the scutellum ; two transverse short ridges placed near the suture, 
one at the middle, the other below the latter and connected by an 
oblique ridge whicli runs from the basal tubercle to a little distance 
from the apex ; another, shorter, oblique ridge extends from the 
shoulder to the middle joining the subsutural one ; a tubercle 
is placed near the apex close to the lateral margin, the suture is 
serrate through its entire length ; the underside rugosely punctate. 

This little species, of which only a single specimen is before me, 



70 MR. M. JACOBY ON THE [Feb. 1, 

differs from E. malayana, Baly, in having more elytral tubercles 
and two, not three, very obsolete longitudinal ridges, the one com- 
mencing at the shoulder joins the subsutural one, with which it 
forms an angle near the middle of the disk ; the transverse and 
other shaped tubercles at and below the middle are very acutely 
raised and form sharp projections. 

Demotina thoracica, sp. nov. 

Greyish fuscous, closely pubescent ; the apices of the tibiae ful- 
vous ; thorax strongly rounded at the middle, obscurely marked 
with brownish bands ; elytra covered with light grey pubescence, a 
spot at the base and two rows of similar spots below the middle, 
placed transversely, fuscous. 

Length H-2 lines. 

Head closely punctured, covered with light grey pubescence, 
which is interrupted at the vertex by two more or less distinct longi- 
tudinal brownish bands ; the anterior margin of the clypeus and the 
labrum fulvous, glabrous. Antennae half the length of the body, 
slender, the terminal joints slightly thickened, the third and fourth 
joints equal, all the joints piceous or dark fuscous. Thorax trans- 
verse, strongly rounded and \Aidened in the middle, the entire sur- 
face covered with whitish-grey pubescence, which assumes the shape 
of a longitudinal band at the sides ; the latter with a round depres- 
sion. Scutellum greyish pubescent. Elytra finely punctate-striate, 
clothed with greyish pubescence like the thorax ; at the basal margin 
two fuscous or dark brown spots are seen, more or less distinct, and 
bounded at the sides by whitish bands ; below the middle tw) trans- 
verse rows of similar spots are placed ; all the femora are armed with 
a strong tooth ; the sides of the elytra are furnished with short and 
stiff bristles. 

Iladley, in Dikoya. 

The thorax in the present species is much narrowed in front and 
at the base, giving more prominence to the middle ; the pubescence 
of the upper surface is generally whitish grey, but sometimes of a 
more fulvous tint, and the fuscous sj)0ts are more or less distinct ; in 
well-marked specimens they are often preceded by whitish sj)ots, 
caused by the more thick pubescence in those places ; in some 
specimens three obHque rows of obscure spots may be seen at each 
elytron besides those at the base, which are generally present. 

Demotina semifasciata, sp. nov. (Plate X. fig. I.) 

Obscurely fuscous or piceous ; finely pubescent ; antennae and 
legs dark fulvous ; scutellum white ; elytra closely and distinctly 
semipunctate-striate, with an obscure darkish transverse band below 
the middle ; a spot at the base and four or five others below the 
middle, white ; femora strongly toothed. 

Length 2-3 lines. 

Head clothed with yellowish pubescence at the vertex, which hides 
the punctuation ; e|)istome imj)ubescent, distinctly punctured, ful- 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 71 

vous ; eyes entire ; palpi slender. Antennae filiform, fulvous, the 
third and fourth joints slender, nearly equal, the terminal joints very 
slightly thickened. Thorax twice as broad as long, the lateral 
margin distinct, the anterior angles produced into a short tooth ; 
surface covered with yellowish, slightly curved hairs like the head. 
Scutellum pentagonal, clothed with thick white pubescence, margined 
with piceous. Elytra wider than the thorax ; closely and distinctly 
punctate-striate and pubescent like the thorax, between the shoul- 
ders and the scutellum at the basal margin a white spot is placed, 
four or five others limit the obscure dark transverse band below the 
middle. Underside thickly covered with white scale-like pubescence. 
All the femora armed with a strong tooth ; intermediate tibiae 
emargiuate at the apices ; claws bifid. 

Galle, Balangoda. 

The shape and colour of this species are subject to considerable 
variation, some specimens being much more robust and broader than 
others. The elytral obscure band is just visible with the naked eye 
in most instances, and the spots which hmit it above and below are 
variable in number, white or yellowish, and composed of close and 
thick pubescence ; there are generally three placed above, and two 
below the elytral band. It is possible that Heteraspis albostriata, 
Motsch., may refer to this species, but the description of this author 
is too vague to recognize the species with certainty. 

Demotina lewisi, sp. nov. 

Fuscous or dark piceous, covered with yellowish scale-like |)ube- 
scence ; basal joints of the autennse fulvous ; scutellum whitish ; 
elytra closely punctate-striate, each elytron with two more or less 
distinct rows of white spots. 

Length 2 lines. 

Antennae more than two thirds the length of the bod}^ the third 
and fourth joints equal, the five terminal joints slightly thickened. 
Thorax twice as broad as long, the sides strongly rounded, the 
surface closely and finely rugose-punctate like the bead. Scutellum 
tiiickly clothed with whitish pubescence. Elytra closely covered 
with yellowish scale-like pubescence, the punctuation distinct, close 
and arranged in rows ; eacli elytron with two stripes of whitish 
pubescent spots, of which one is placed at a little distance and close 
to the suture, the other at the middle of the disk and extending 
from the shoulder to the apex ; besides the pubescence single short 
black and stiff bristles are seen on the surface of the elytra. Legs 
dark fulvous, the femora with a distinct tooth. 

Galle. 

i>. lewisi is smaller than the preceding species, the thorax is not 
depressed at the sides and without any stripes or other marks as iu 
D. thoracica ; the elytra are differently marked and without the 
transverse rows of spots as in the last-named species ; the punctua- 
tion of the elytra in D. lewisi is also much deeper and stronger : from 
D. semifasciata the absence of the elytral band sufficiently separates 
the present species. 



72 MR, M. JACOBY ON THE [Feb. 1, 

Demotina ceylonensis, sp. nov. 

Broadly ovate, robust ; fuscous, clothed with fulvous pubescence ; 
the three or four lower joints of the antennae fulvous ; eljtra with 
a transverse whitish band at the middle, the apical portion spotted. 

Length 2 lines. 

Galle. 

Broader and more robust than D. thoracica, and the pubescence 
fulvous instead of whitish. The elytra, instead of finely and distantly 
punctate-striate, are here strongly and closely punctured, and a 
transverse hand, comjiosed of thick white pubescence, is placed at 
the middle ; this band is narrowed towards the suture and is, in one 
specimen, followed by a broad black denuded space, while the apical 
portion is variegated by white and fulvous pubescence ; in another 
specimen, which I refer to the same species, the transverse band is 
only indicated and the space below it shows some small fuscous 
spots, alternated by white and fulvous pubescence. The present 
species resembles somewhat D. fasciata, Baly, but is more robust 
and the thorax is less transverse, the pubescence shorter and differ- 
ently placed. 

Xanthonia flavopilosa, sp. nov. 

Narrowly elongate ; pale fulvous, covered with fine silky flavous 
pubescence ; terminal joints of the antennae dusky ; elytra extremely 
minutely punctured. 

Length l|-2 lines. 

Head extremely finely punctured, covered with rather long 
yellowish hairs ; the anterior margin of the epistome nearly straight ; 
eyes entire, scarcely sinuate within. Antennae nearly as long as the 
body in the male, the third and fourth joints nearly equal. Thorax 
one half broader than long, transversely depressed across the disk, 
the sides strongly rounded, the lateral margin obsolete ; the surface 
clothed, like the elytra, with rather long silky yellow pubescence, 
extremely finely punctured ; femora unarmed ; tibite entire ; claws 
bifid ; the anterior margin of the thoracic cpisternum subconcave. 

Galle, Dikoya. 

This species is larger than X. placida, Baly, from Japan ; the 
thorax is more transverse and flattened, and the punctuation of the 
upper surface is much more finely impressed, and only visible under 
a strong lens ; the pubescence is also longer. 

Nephrella elongata, Baly. 

I do not think I am wrong in referring the two specimens contained 
in this collection to Baly's species, witli the description of which they 
agree perfectly, except in one respect in regard to the punctuation 
of the upper surface. In the specimens before me the thorax and 
elytra, which are of an obscure fuscous, are closely covered with fine 
fulvous pubesence, which totally obscures any punctuation. Mr. Baly 
speaks of the thorax as not very deeply punctured, and of the elytra 
as finely wrinkled, of which I am not able to see a trace. It is 
therefore possible that the specimens obtained by Mr. Lewis repre- 



188/.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 73 

sent a distinct species ; but as they closely agree in all the other cha- 
racters pointed out by that author, I have abstained from describing 
them as new. The type of Nephrella seems unfortunately to have 
been lost, as it is not contained in Mr. Baly's collection novr in the 
British Museum. I may add, further, that the head and thorax in 
Mr. Lewis's specimens show a fine central raised ridge, of which 
Mr. Baly says nothing. 

Chrysolampra punctatissima, sp. nov. 

^neous ; antennae and legs piceous ; head and thorax extremely 
closely and finely punctured ; elytra strongly transversely strigose, 
finely punctured near the suture only. 

Var. Dark purplish blue, subopaque. 

Length 3-4 lines. 

Head extremely finely and closely punctured, the anterior margin 
of the clypeus emarginate at the sides and middle ; labrum fulvous. 
Antennae slender, the terminal joints very slightly thickened, the two 
basal joints generally fulvous, the rest piceous. Thorax twice as 
broad as long, the sides rounded, tuberculate at the anterior angles ; 
surface rather convex, as closely and a little more distinctly punc- 
tured than the head, the interstices slightly rugose at the sides ; 
scutellum dark purplish, impunctate, as broad as long. Elytra sub- 
cylindrical, the entire disk covered with strongly raised transverse 
rugosities, which near the apices form single tubercles ; the space 
near the base and at the suture remotely and finely punctured, the 
latter accompanied near the apex by one or two longitudinal costse. 
Legs piceous or dark fulvous ; the anterior femora dilated at the 
middle and with a more or less distinct tooth ; anterior thoracic 
episternum concave. 

Galle. 

Principally distinguished by the very close punctuation of the head 
and thorax. 

Pagria costatipennis, sp. nov. (Plate X. fig, 2.) 

Subquadrate-ovate ; bronze-coloured; three or four basal joints of 
the antennae fulvous ; head rugose-punctate ; thorax longitudinally 
strigose and deeply punctured, subcylindrical, convex ; elytra longi- 
tudinally costate, the interstices deeply punctured. 

Length l|-li line. 

Head coarsely rugose- punctate, deeply sulcate above the eyes; 
anterior margin of the epistome concave-emarginate ; labrum and 
palpi fulvous, the hitter slender. Antennae more than half the length 
of the body in the male, shorter in the female ; the third and fourth 
joints slender, of equal length and about one half longer than the second 
joint, the terminal joints obscure piceous. Thorax twice as broad as 
long, the disk strongly swollen, the sides deflexed anteriorly, the 
entire surface closely and strongly elevate, reticulate and strigose, 
the interstices forming deep punctures and foveas. Scutellum sub- 
pentagonal, its surface longitudinally depressed. Elytra subquadrate- 
ovate, broader at the base than the thorax, with a distinct depression 



M- 



74 MR. M. JACOBY ON THE [Feb. 1, 

below the base, which interrupts the longitudinal costae, which latter 
are entire from there to the apices, those at the sides being frequently 
broken and shorter, and the interstices coarsely rugose and wrinkled. 
The legs are piceous, the apices of the tibiae and the tarsi obscure 
dark fulvous. 

Galle. 

A closely allied but distinct species belonging to this genus has 
been described by Walker in the 'Annals and Mag. of Nat. Hist.' for 
1858 as a Curculio and a Rhijnchites, with which it has of couise 
nothing in common ; that species, I believe also from Ceylon, is 
contained in the collection of the British Museum. I have placed 
the present species in M. Lefevre's genus Payria (Bull, de France, 
1884), on account of the sulcation above the eyes, although the 
convex shape of the thorax and its rounded sides differ from that of 
the species described by M. Lefevre ; but as theangnlate and rounded 
margins of the thorax are both met with in the genus Nodostoma, it 
would not be wise to establish another genus on that character only, 
the more so as all other structural characters peculiar to Pagria are 
present in the species here described. 

NODINA SUBDILATATA, Motsch. 

A specimen named as above and contained in the collection of 
Mr. Baly agrees with those obtained by Mr. Lewi^. The description 
given by Motschulsky is too superficial and almost useless, and it is 
therefore on the authority of Mr. Baly that I refer the present 
species to that of Motschulsky. N. su/jdilatata seems to me to 
possess but little affinities in regard to structural characters to those 
species described subsequently b}^ Mr. Baly and placed in Nodina. 
Pagria, Lefcv., seems the proper genus in which to place the present 
species, as it agrees in general shape, the armed femora, and the 
orbital grooves, as well as in most other details, with that genus. I 
give here a renewed description of the species before nie. 

Subquadrate-ovate ; black, above metallic green or seiieons ; basal 
joints of the antennee, the base of the posterior femora, and the 
tibiae and tarsi fulvous ; thorax closely and deeply punctured ; 
elytra with basal depression, strongly punctate-striate. 

Length 1-1 1 line. 

Head deeply but not closely punctured, deeply sulcate above the 
eyes ; epistome more closely punctured ; labrum and palpi fulvous. 
Antennae more than half the length of the body, the four basal 
joints fulvous, the rest l)lack ; the second joint thickened, the third 
and fourth joints slender, of equal length, the others slightly 
thickened. Thorax one half broader than long, subcylindrical, con- 
vex, the sides rounded ; the surface closely covered with deep and 
round punctures, slightly transversely sulcate near the anterior 
margin. Elytra with a distinct depression below the base, very 
deeply punctate-striate, the interstices at the sides and near the 
apices costate, the punctuation much less deeply impressed below 
the middle ; femora dark aeneous, their apices more or less fulvous, 
tibiae and tarsi light fulvous ; the four posterior tibiae emarginate 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 75 

near the apices ; claws appendiculate ; femora with a very minute 
tooth ; prosternum broader than long, strongly punctured. 

Variable in size and colour and allied in regard to the latter to 
Nodostoma jansoin, Baly, and N. tibiale ; the distinct elytral depres- 
sion will separate the species from the former, and the very closely 
punctured thorax from the last-named species. It is quite possible 
that Noda viridicenea, Sclionh., refers to the present species. 

Rhyparida L^vicoLLis, sp. nov. 

Obscure fulvous ; eyes closely approached ; head and thorax 
impunctate ; elytra with a subbasilar depression, strongly punctate- 
striate ; femora dilated into a strong triangular tooth. 

Length 2 lines. 

Head impunctate ; eyes very large, the space dividing them 
narrower than their diameter, their inner margin deeply notched ; 
epistome separated from the front by a few punctures only.' Antennae 
nearly as long as the body, fulvous, the fourth joint longer than the 
third, this longer than the second joint, the following slightly 
thickened, the apices of the joints stained with fuscous. Thorax 
transverse, the sides rounded, the angles distinct, the surface entirely 
impunctate or with a few microscopically fine punctures. Elvtra 
with a distinct depression below the base, the latter somewhat 
swollen, the surface deeply punctate-striate at the anterior portion, 
more finely punctured towards the apices; the anterior and posterior 
femora dilated into a strong triangular tooth, the intermediate 
femora minutely dentate ; claws bifid. 

Galle. 

R. Icpvicollis may be recognized by the large and closely approached 
eyes and the impunctate thorax, in connection with the strongly 
dentate femora ; the elytra have the basal portion more distantly 
punctured than the rest of the surface, and the shoulders are entirely 
devoid of punctuation, but bounded within by a deep hue of closely 
approached punctures, the latter being more widely placed below 
the elytral depression ; the general colour varies from pale to dark 
fulvous, the sides and the suture being sometimes stained with 
piceous. No species of RJnjparida has, to my knowledge, hitherto 
been recorded from Ceylon. 

Rhyparida quinquemaculata, sp. nov. (Plate X. fig. 3.) 

Rufous ; the last eight joints of the antennae and the legs black ; 
thorax sparingly and finely punctured ; elytra regularly punctate- 
striate, a sutural spot at the middle, another at the shoulder, and a 
third near the apex of each elytron black ; femora toothed. 

Length 2 lines. 

Head with a deep fovea at the vertex, not visibly punctured ; the 
epistome separated from the face by a slight transverse depression. 
Antennae half the length of the body, black, the three lower joints 
fulvous, the third and fourth joints slender, of equal length, the 
following joints slightly depressed and shorter. Thorax transversely 
convex, about three times as broad as long, the sides rounded ; the 



7b MR. M. JACOBY ON THE [Feb. 1, 

surface very finely and sparingly punctured, rufous, with a small 
obscure piceous spot at the sides. Elytra with a distinct transverse 
depression below the base, the punctured striae very distinct 
anteriorly, less so posteriorly ; each elytron with a square-shaped 
black spot at the shoulder, a triangular one near the apex, and a 
common sutural elongate spot near the middle ; sides of the breast 
and the legs black ; all the femora with a small tooth ; claws bifid. 

Dikoya. 

A single specimen. 

NODOSTOMA BITUBERCULATUM, Sp. nOV. 

Testaceous ; the terminal joints of the antennae black ; thorax 
strongly and closely punctured, angulate below the middle ; elytra 
strongly punctate-striate anteriorly, each elytron with two tubercles 
placed at the shoulders. 

Length 1 line. 

Head very strongly and remotely punctured ; the epistome not 
separated from the front. Antennae nearly as long as the body ; the 
second joint not much shorter than the first, the fourth joint slightly 
longer than the third. Thorax twice as broad as long, narrowed in 
front, the sides angulate near the base ; the surface strongly and 
closely rugose-punctate, without an anterior transverse groove. 
Elytra very slightly depressed below the base, the punctuation 
almost entirely absent near the apices ; the humeral callus in shape 
of an elongate tubercle, which is followed immediately below by 
another smaller tubercle, the space between these latter and the 
lateral margins deeply depressed ; underside of a more fulvous tint ; 
femora with a minute tooth. 

Dikoya. 

The small size, closely and strongly rugose thorax, and the lateral 
tubercles of the elytra will help to separate the present species from 
its many congeners. 

NoDOSTOMA BIPUNCTATUM, Sp. UOV. 

Testaceous ; thorax angulate at the sides, finely punctured ; 
elytra indistinctly punctured below the middle, the sutural and 
lateral margins and a spot below the base on each elytron black. 

Var. Elytra entirely black. 

Length 2 lines. 

Head with a few fine punctures between the eyes ; the epistome 
not separated from the face ; eyes distinctly sinuate. Antennse 
slender and filiform, the apical joints not thickened, testaceous, the 
fourth joint distinctly longer than the third. Thorax short, narrowly 
transverse, greatly widened towards the base, the sides subangulate 
close to the latter ; the surface with a narrow transverse groove in 
front of the anterior margin, rather closely and finely punctured. 
Scutellum obscure fulvous or pioeous ; its apex broadly rounded. 
Elytra with a deep depression below the base, distinctly punctured 
above this depression only, the rest of the surface obsoletely punctate, 
the interstices very slightly raised : the sutural and lateral margins 



1887.] PHYTOPHAGOUS COLEOPTKRA OF CEYLON, ^^ 

narrowly black, the latter accompanied by a row of deep punctures. 
Legs testaceous; all the femora armed with a small toolh. 

Dikoya. 

Amongst the many described species o^ Nodostoma, N. bipunctatum 
may be known by the very short and transverse thorax in connection 
with the coloration. 

NODOSTOMA IMPRESSIPENNE, Sp. noV. 

Entirely testaceous ; the last seven joints of the antenna black ; 
head and thorax distantly punctured ; elytra with a deep basal 
depression, the latter strongly, the rest finely punctate-striate ; legs 
very long. 

Length 1| line. 

Head strongly but very remotely punctured ; the epistome not 
separated from tlie face. Antennae slender, scarcely shorter than the 
body, the four lower joints testaceous, the rest black, the third and 
fourth joints slender, equal. Thorax not more than twice as broad 
as long, the sides angulate near the base, obhquely narrowed 
towards the apex and slightly rounded before the middle ; the surface 
with a transverse groove in front of the anterior margin, very 
strongly but remotely punctured. Elytra narrowed posteriorly, with 
a deep transverse depression below the ba.se, testaceous, the sutural 
and lateral margin narrowly fulvous; the punctuation deep within 
the depression, very fine at the rest of the surface ; there is also a 
row of deep punctures placed close to the lateral margin and below 
the shoulders ; underside and the legs testaceous ; all the femora 
armed with a small tooth. 

Galle. 

Closely allied to N. fairmairei, but the thorax is longer, less 
transverse, and more strongly and remotely punctured, and the legs 
are much longer in N. impressipenne, 

NoDOSTOMA LEWISI, Sp. HOV. 

Testaceous or fulvous ; the apical joints of the antennee black ; 
head remotely, thorax very closely and strongly punctured, sub- 
angulate near the base ; elytra nearly impunctate below the middle, 
the sutural and the lateral margins black. 

Var. The disk of the thorax more or less piceous. 

Length 1-1^ line. 

Head with a few punctures ; the epistome more strongly and 
closely punctured, not separated from the front. Antennse two 
thirdsthe length of the body, the four lower and the base of the 
followmg jomts testaceous, the terminal joints black. Thorax 
transverse, strongly narrowed in front, the sides distinctly angulate 
near the base, the surface very strongly or subrugosely jpunctured. 
Elytra with a distinct depression directly below the base, the latter 
distmctly punctured, the punctuation gradually disappearing below 
the depression; the lateral margin narrowly, the sutural more 
broadly, black, this colour widened in some specimens towards the 
base at the suture. Legs with a very minute tooth. 

Dikoya. 



78 MR. M. JACOBY ON THE [Feb.], 

N. lewisi resembles somewhat N. consimile, Baly, from Jap m ; but 
that species has shorter and eiitirel}' fulvous auteuuae, the thorax is 
less transverse and less strongly punctured, while the elytra have 
the punctures deeper and more prolonged posteriorly. Whether 
N. triangulare, Motsch., also from Ceylon, is referable to the species 
described here it is impossible to say, on account of tlie short and 
insufficient description. The black sutural stripe is in some specimens 
very narrow, in others widened towards the suture. 

NoDOSTOMA TUBEROSUM, sp. nov. (Plate X. fig. 4.) 

Dark brown ; head closely punctured ; thorax strongly rugose- 
punctate, the sides angnlate near the base ; elytra entirely covered 
with longitudinal and transverse tuberosities. 

Length 3 lines. 

Head closely and distinctly, the vertex more remotely, punctured ; 
epistome not separated from the face, its anterior margin perfectly 
straight ; the surface covered with some short silvery pubescence. 
Antennae with the first three joints fulvous (the rest wanting). 
Thorax transverse, narrowed in front, the sides distinctly angulate 
near the base ; the surface entirely covered with deep and round 
punctures, the interstices sparingly clothed with short hairs. Elytra 
closely covered with strongly raised tubercles, placed irregularly at the 
sides, but arranged in longitudinal rows at the disk ; the interstices 
with some deep punctures ; the shoulders prominent and in the shape 
of an oblique, smooth, strongly raised tubercle. 

A sinde specimen is contained in my collection. N. tuberosum is 
not difficult to recognize, on account of the wart-like tubercles 
covering the entire surface of the elytra. 

NoDOSTOMA LEFEVREl, Sp. nOV. 

Dark fulvo-piceous, the base of the femora fulvous ; thorax 
transverse, angulated at the base, very closely punctured; elytra 
-with deep basal depression, strongly longitudinally costate, the sides 
transversely rugose. 

Length 2g lines. 

Head closely punctured at the vertex and at the sides, the 
epistome separated from the face by two deep fovese at the sides, its 
anterior edge tridentate, the surface scarcely visibly punctured ; 
labrum fulvous. Antennae slender, the third and following joints 
elono-ate. Thorax very transverse, three times broader than long ; 
the sides strongly angulate near the base, the posterior angles pro- 
duced into an acute tooth ; the surface with a transverse groove near 
the anterior margin, very closely punctured, with some smooth and 
raised spaces irregularly distributed. Scutellum nearly black, 
impunctate. Elytra with a deep basal depression, very strongly 
costate at the inner portion of the disk, the interstices regularly 
punctate-striate, the sides transversely rugose and wrinkled; the 
colour fighter fulvous near the suture, darker at the sides ; femora 
with a minute tooth, their base pale fulvous. 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 79 

A single specimen (coll. Jacoby). 

N. lefevrei differs from N. tuberosum in the much more transversely- 
shaped and finely punctured thorax, the strongly costate elytra and 
their coloration. 

NODOSTOMA CLYPEATUM, Sp. nOV. 

Pale testaceous; apical joints of the antennae fuscous; clypeus 
strongly punctured ; thorax angulate below the middle, strongly 
punctured at the sides only ; elytra with basal depression, the latter 
distinctly, rest of the disk nearly obsoletely, punctured. 

Length 1 line. 

Head nearly impunctate at the vertex ; the epistome strongly but 
very remotely [)unctured, not separated from the front ; the space 
above the insertion of the antennas obliquely grooved. Antennae 
slender, nearly as long as the body, testaceous, the three or four 
terminal joints fuscous or black. Thorax twice as broad as long, 
strongly narrowed in front, the sides distinctly angulate near the 
base, surface without an anterior groove, strongly punctured at the 
sides only, the middle of the disk with a iev; fine punctures only. 
Elytra with a very distinct subbasilar depression, the base itself with 
a few remotely placed punctures arranged in hues which extend 
more or less distinctly to the middle; below the latter the punc- 
tuation is almost entirely wanting ; femora with a minute snine. 

Galle. ^ 

Principally distinguished by the nearly impunctate vertex, the 
strongly punctured epistome and sides of the thorax, in connection 
with its general small size ; the absence of a transverse anterior 
groove at the thorax will further assist in the recognition of 
iVi clypeatum. 

NoDOSTOMA LONGICORNE, Sp. nOV. 

Fulvous ; legs testaceous, the knees obscure piceous ; antennae 
as long as the body ; thorax very finely punctured, angulate behind 
the middle ; elytra strongly punctate-striate, with basal depression, 
the lateral margins anteriorly black. 

Var. Elytra entirely fulvous. 

Length 2-2g lines. 

Head finely and rather remotely punctured ; the epistome separated 
from the front by a slight transverse depression, with a few deep 
punctures ; palpi testaceous. Antennae as long as, or slightly lono-er 
than, the body, fulvous, the apical joint darker, the fourth joint rather 
longer than the third. Thorax transverse, strongly narrowed in front, 
the sides distinctly angulate near the base, the surface with a deep and 
punctured transverse sulcation near the anterior margin, very finely 
and ratiier remotely punctured, the lateral margin narrowly piceous. 
Elytra not broader at the base than the thorax, with a deep subbasal 
depression, very strongly and deeply punctate-striate anteriorly, more 
finely towards the apices, the interstices slightly convex, more dis- 
tinctly so at the sides, the lateral margin anteriorly rather broadly 
piceous or black, this colour extending slightly across the elytral 



80 MR. M..TACOBY ON THE [Feb. 1, 

depression, but being narrowed posteriorly along the lateral margin. 
Legs long and slender, testaceous, the knees obscure piceous ; the 
femora armed with a small tooth. 

The variety, which I believe is a female, is of larger size, the 
elytra are less strongly punctured and entirely dark fulvous ; the 
terminal joints of the antennae are also stained with fuscous ; but in 
all other respects this specimen agrees with the type. N. longicorne 
may be distinguished from numerous other similarly coloured species 
by the long antennae and the finely punctured head and thorax. 

NODOSTOMA FAIRMAIREI, Sp. nOV. 

Pale testaceous ; antennae (the two or three basal joints excepted) 
black, tarsi stained with piceous ; thorax strongly and remotely 
punctured ; elytra with basal depression, strongly punctured ante- 
riorly, more finely towards the apices. 

Length l|-2 lines. 

Head remotely but strongly punctured, the epistome not separated 
from the face. Antennae two thirds the length of the body, the 
first joint short and dilated, the third and fourth thin and of nearly 
equal length, the others slightly thicker. Thorax scarcely twice as 
broad as long in the male, the sides subangnlate below the middle, 
narrowed in front, the surface strongly punctured at the sides and 
anteriorly, the interstices slightly rugose. Elytra subcyliudrical, 
parallel, distinctly depressed below the base, the punctuation rather 
strong anteriorly, gradually diminishing posteriorly. Legs rather 
long ; the femora armed with a small tooth ; the extreme apices of 
the tibife and the tarsi stained with piceous. 

Dikoya. 

N.fairmairei differs from N. ohliteratum in the much stronger 
punctuation of the head and thorax, the shorter and nearly black 
antennae, and in the similarly coloured apices of the tibae. 

NODOSTOMA OBLITERATUM, Sp. UOV. 

Pale flavous ; antennae as long as the body ; head and thorax 
with a few fine punctures, the latter angulate near the base ; elytra 
with basal depression, very finely punctured anteriorly, the punctures 
nearly obsolete below the middle. 

Length 11-2 lines. 

Head with a few scarcely visible punctures ; the epistome not 
separated from the front ; jaws piceous. Antennte slender, fulvous, 
the terminal joints more or less stained with fuscous. Thorax nearly 
three times as broad as long, the sides strongly narrowed in front, 
distinctly angular near the base ; surface with a distinct narrow 
groove close to the anterior margin, very finely and sparingly punc- 
tured. Elytra slightly broader at the base than the thorax, the 
sides parallel, the shoulders moderately prominent, the subbasilar 
depression distinct but not deep, the punctuation much more dis- 
tinctly visible anteriorly than posteriorly, where only traces of it can 
be seen under a strong lens ; femora armed with a small tooth. 

Dikoya. 



1887.] PHYTOI'HAGOUS COLKOPTERA OF CEYLON. "81 

Dermorrhytis cuprea, sp. nov. 

Bronze-coloured ; antennae and legs nearly black ; thorax sub- 
remotely and strongly punctured, margined with green ; elytra 
distantly punctate-striate, the sides transversely rugose below the 
shoulders. 

Length 3-3| lines. 

Head strongly rugosely punctured at the vertex ; the epistome 
niuch more closely punctured and separated from the face, its lower 
edge straight ; labrum and jaws piceous. Antennae filiform, two 
thirds the length of the body, the third and three following joints 
piceous, slender, and of equal length, the others slightly thickened 
and nearly black. Thorax not more tlian twice as broad as long, 
the sides obsoletely angulate before the middle, nearly straight from 
there to the base, the margin slightly dentate or sinuate, metallic 
greenish ; rest of surface strongly and remotely punctured, the punc- 
tuation more close and strong at the sides, where the interstices are 
partly rugose and forming transverse wrinkles. Elytra cupreous, 
with more or less metallic greenish reflections, very distantly and 
strongly punctate-striate at the disk, the sides very strongly trans- 
versly rugose and wrinkled. Prosternum dilated posteriori v, broad ; 
the anterior margin of the thoracic episternum concave. 

D. cuprea may be recognized by the distant punctuation of the 
elytra, the general cupreous colour of the upper surface, and the 
nearly black legs. The punctuation of the thorax differs according 
to the sex, and is sometimes more closely arranged ; while in some 
specimens the middle of the disk represents a nearly smooth longi- 
tudinal space ; the anterior angles are rather prominent, and the 
shape of the thorax is less transverse than in the other allied forms. 

Dermorrhytis ceylonensis, sp. nov. (Plate X. fig. 7.) 

Greenish or brownish aeneous, below obscure piceous ; antennsD 
and legs fulvous ; head and thorax closely and rather finely rugose- 
punctate ; elytra punctate-striate, the interstices anteriorly and at 
the sides transversely strigose. 

Length 21-3 lines. 

Head closely rugose-punctate, the space between the antennae fur- 
nished with a smooth tubercle ; labrum fulvous. Antennae slender, 
two thirds the length of the body, fulvous, the terminal joints some- 
times stained with piceous, tliird joint slightly longer than the fourth. 
Thorax scarcely twice as broad as long in the male sex, more trans- 
verse in the female, the sides very slightly rounded or obsoletely 
angulate before the middle, the anterior angles acute and slightly 
produced ; surface closely rugose-punctate like the head, metallic 
green, the extreme lateral margin reddish cupreous anteriorly. 
Elytra much more remotely and more strongly punctured than the 
thorax, the interstices strongly raised aud forming transverse strig© 
anteriorly and at the sides, as well as at the apices, the lateral 
margin bright aureous-cupreous. Legs dark fulvous or obscure 
piceous. 

Dikoya. 
Proc. Zool. Soc— 1887, No. VI. 6 



S2 MR. M. JACOBY ON THE [Feb. 1, 

Easily distinguished by tlie densely rugose-punctate thorax and 
the cupreous margin of the latter and of the elytra. 

Dermorrhytis lewisi, sp. nov. 

Greenish or brownish cupreous ; basal joints of the antennae and 
the legs obscure fulvous ; head and thorax closely and strongly 
punctured ; elytra strongly punctate-striate, the interstices trans- 
versly strigose throughout. 

Length 2|-3 lines. 

Head closely rugose-punctate, the epistome not separated from 
the face ; labrum fulvous. Antennae very slightly thickened at 
the apical joints, the third and fourth joints equal, the five or six 
lower joints fulvous, the others fuscous. Thorax transverse, the 
sides nearlj' straight, the surface closely and strongly punctured, 
the interstices everywhere transversly rugose. Elytra subcylindrical, 
slightly narrowed behind, punctured like the thorax, but the inter- 
stices transversly raised and extending across the entire surface ; 
underside covered with fine silky-white pubescence. 

Dikoya. 

The straight sides of the thorax, in connection with the trans- 
versely rugose interspaces of the same part, and those of the entire 
disk of the elytia, will help to distinguish this species principally 
from B. piceipes, Brdy. 

Dermorrhytis ornatissima, Baly. (Plate X. fig. 6.) 

Dermorrhytis fasciato-rutilans, Leto vre. 

Balangoda. 

This species, of which the two above names are synonymous, seems 
to vary greatly in regard to size and coloration ; all the specimens 
contained in my collection are devoid of any cupreous spots on the 
thorax. In the present collection the specimens obtained by Mr. 
Lewis are much smaller, and have a broad transverse reddish-cupreous 
band occupying nearly the entire disk of the thorax. 

Dermorrhytis variabilis, sp. nov. 

Greenish aeneous below ; above metallic green, the shoulders, 
sides, and apices of the elytra reddish cupreous ; head and thorax 
remotely punctured, the interstices flat ; elytra punctate-striate, 
the sides transversely strigose ; basal joints of the antennae and the 
legs fulvous. 

Var, Above brownish cupreous. 

Length 2-2| lines. 

Head rather flattened, finely and moderately closely punctured, 
the interstices slightly rugose anteriorly only ; the epistome not 
separated from the face, its anterior margin nearly straight ; labrum 
fulvous. Antennte longer than half the length of the body, the five 
terminal joints slightly dilated, black, the others fulvous. Thorax 
nearly twice as broad as long, the sides straight and slightly subangulate 
or dentate; surface remotely and distinctly punctured, the interstices 
not raised. Elytra punctate-striate at the inner disk, irregularly 



188/.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 83 

and more strongly punctured at the sides, where the interstices are 
transversely and strongly raised ; the apex of each elytron is simply 
punctate-striate and the interstices are flat. 

Galle. 

The smaller size, remotely punctured thorax, and the colour of 
the upper parts distinguish this species from its allies. 

Dermorrhytis igneofasciata, Baly. (Plate X. fig. 5.) 

Melasoma divisi, sp. nov. 

Fulvous ; terminal joints of the antennse and the scutellum black ; 
elytra distinctly punctured, testaceous, the suture and a longitudinal 
band on each elytron, divided at its posterior portion, metallic 
greenish. 

Length 2 lines. 

Head extremely finely punctured, longitudinally depressed at the 
middle. Antennae short, the six terminal joints transversely shaped 
and black, the others fulvous. Thorax more than three times 
broader than long, slightly narrowed iu front, the sides but little 
rounded, the posterior margin broadly rounded at the middle and 
slightly produced ; the surface with a few minute punctures at the 
sides. Scutellum black. Elytra rather strongly and closely punc- 
tured, the shoulders prominent, the lateral margin slightly thickened 
and the space in front impunctate ; at the middle of the disk a 
broad metallic green band is placed, commencing at the base and 
extending with its inner division below the middle, tiie outer portion 
being prolonged in a narrow stripe to the apex of each elytron, where 
it is joined to the similarly coloured sutural margin. Legs more or 
less stained with piceous. 

A single specimen. 

Haltica (Graptodera) (?) NiGRiPENNis, sp. nov. 

Flavous ; thorax transverse, impunctate, the transverse groove 
not extending to the sides ; elytra black, extremely finely and 
closely punctured. 

Var. Antennae and legs piceous. 

liength 2 lines. 

Head impunctate, the frontal tubercles obsolete, scarcely raised. 
Antennae half the length of the body, the second joint very short, 
the five following joints slightly triangularly shaped and widened, 
the rest more elongate. Thorax twice as broad as long, the sides 
rather rounded at the middle, the angles obsolete, surface entirely 
impunctate, impressed near the base with a shallow transverse 
groove not extending to the sides. Scutellum flavous. Elytra 
nearly parallel, very finely punctured (in one specimen impunctate), 
black ; the first joint of the posterior tibiae as long as the two 
following joints together ; claws appendiculate ; the anterior coxal 
cavities open. 

The present species has certainly not many of the typical cha- 
racters peculiar to Graptodera ; the more transverse thorax, the 

6* 



84 MR. M. JACOBY ON THE [^''^b. 1, 

different shape of its groove, and the general colour seem to point 
to an allied but different genus. 

Enneamera ceylonensis, sp. nov^. 

Testaceous ; a spot at the vertex and the soutellum black ; head 
and thorax inipunctate ; elytra scarcely visibly punctured, testa- 
ceous, a triangular-shaped spot at the base and a narrow trans- 
verse hand below the middle reddish fulvous. 

Var. Elytra entirely dark fulvous. 

Length 2 lines. 

Head broader than long, impunctate, a large spot at the vertex 
black ; frontal tubercles obsolete ; labrum piceous. Antennae 
entirely pale fulvous, the second and third joints short, the rest 
transversely dilated. Thorax three times as broad as long, widened 
at the middle, the sides nearly straight, narrowed in front, the 
anterior angles slightly thickened and distinct ; the surface entirely 
impunctate, pale testaceous. Scutellum black. Elytra microscopi- 
cally finely punctured, very convex and rounded, of a yellowish 
colour, a transverse triangular-shaped band at the base, not quite 
extending to the sides, and marked with a more or less distinct black 
spot at the shoulder, reddish fulvous ; the narrow band behind the 
middle of a more piceous colour ; the outer margin of the posterior 
tiliise with a row of small blaclv teeth or spines. 

Besides the above-named fulvous variety, which does not vary in 
other respects from the typical form, I possess a specimen in my 
collection (also from Ceylon) in which the posterior band of the 
elytra is divided into two small spots. 

Phyllotreta discoidea, sp. nov. 

Head, thorax, and the abdomen fulvous ; antennae, the breast, and 
legs black ; elytra scarcely visibly punctured, testaceous, all the 
margins narrowly black. 

Length 1| line. 

Head impunctate, fulvous, the frontal tubercles transverse, narrow 
and very distinct. Antennae half the length of the body, black, the 
third joint smaller than the second, the following ones gradually 
thickened. Thorax transversely quadrate, the sides slightly rounded ; 
ths surface rather flat, with a very obsolete and shallow depression 
at the middle of the sides, entirely impunctate. Scutellum black. 
Elvtra parallel, not covering the pygidium, their surface only visibly 
punctured when seen under a strong lens, testaceous or yellowish, 
margined with black, the sutural margin generally narrowed near 
the base. Breast and legs black, the anterior femora slightly stained 
with fulvous below ; the abdomen, with the exception of the last 
segment which is black, fulvous. 
Bogawantalawa. 

P. discoidea may be recognized by the small third joint of the 
antennae, which is smaller than the second (an exceptional structure 
as a rule), and by its general coloration. 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 85 

Aphthona ceylonensis, sp. nov. 

Ovate ; obscure piceous ; antennae, tlie apices of the tibise, and the 
tarsi testaceous ; thorax transverse, very finely punctured ; elytra 
more distinctly and very closely punctate. 

Var. Entirely obscure testaceous. 

Length | line. 

Head iinpunctate ; the frontal tubercles narrowly transverse. 
Antennae closely approached, nearly as long as the body, testaceous, 
the terminal joints more or less stained with fuscous, the third and 
the two following joints nearly equal, smaller and thinner than the 
second, the following ones slightly thickened. Thorax much broader 
than long, the sides slightly rounded, the anterior angles oblique ; 
the surface rather closely and finely punctured, the interstices 
extremely finely alutaceous. Elytra distinctly widened at the middle, 
rather convex, the shoulders rounded, closely and very distinctly 
punctured, the interstices somewhat rugose ; the legs piceous, the tibias 
more or less tesiaceous ; the first joint of the posterior tarsi as long 
as the two following together, 

Horton Plains. 

This very small species seems to vary much in regard to colour 
from nearly black to testaceous, and several intermediate degrees are 
before me. The ovate, anteriorly and posteriorly narrowed shape of 
the elytra and their close punctuation, as well as that of the thorax, 
and tlie small size of the insect will help in the recognition of A. 
ceylonensis. 

Aphthona lewisi, sp. nov. 

Fulvous ; antennae with the fifth to the tenth joints black ; thorax 
finely and subremotely punctured; elytra black or piceous, veiy 
closely punctured. 

Length 1 line. 

Head impunctate, shining, fulvous. Antennae two thirds the length 
of the body, the four basal joints fulvous, the following six black, the 
apical joint fulvous. Thorax twice as broad as long, the sides straight, 
slightly converging towards the apex, tiie anterior angles obliquely 
truncate, forming a distinct angle before the middle ; the surface 
covered throughout with very fine but not very closely placed 
punctures. Scutellum obscure fidvous. Elytra nearly parallel, sub- 
cylindrical, the shoulders distinct but not prominent, the surface 
more distinctly and more closely punctured than the thorax ; 
underside and legs fulvous. 

Bogawantalawa. 

A. lewisi may be known by the general coloration, the finely 
punctured thorax, and the colour of the antennae, which agree in all 
the specimens before me. 

Aphthona proxima, sp. nov. 

Obscure piceous ; the five or six basal j(jints of the antennae fulvous ; 



86 MR. M. JACOBY ON THE [Feb. 1, 

thorax finely punctured ; elytra more distinctly and closely punctate, 
the interstices slightly rugose. 

Length 1 line. 

Head impunctate, the frontal tubercles distinctly raised. Antennae 
nearly as long as the bod}', rather robust, the second joint nearly as 
stout as the first, but shorter ; the three following joints more 
slender, of equal length, the following slightly thicker. Thorax 
transverse, the anterior angles oblique, notched before the middle, 
the sides straight, sligiitly converging outwards ; the surface finely 
and rather closely punctured. Elytra with a shallow depression below 
the base, somewhat closely and rugosely punctured, the punctuation 
visible to the apices ; legs short and robust. 

Balangoda. 

J. proximo is extremely closely allied to A. sordida, Baly, from 
Japan, and may possibly be identical with that species ; but the 
depression of the elytra below the base in the Ceylon specimens does 
not permit me to refer them to that species, as I cannot see a similar 
depression in A. sordida, of which I possess several specimens. The 
antennae in the present insect seem to vary rather in colour, and in 
one of the specimens, wliich I look upon as a variety, having been 
taken together with the others, the third and the fifth joints of the 
antennae are fulvous, the others black ; in this specimen the elytral 
depression is also much more marked (the base being slightly raised) 
than in the others. 

Aphthona vicina, sp. nov. 

Ovate, convex ; black ; the third and one or two following joints of 
the antennae flavous ; tliorax finely and remotely punctured ; elytra 
niore distinctly and closely semipunctate-striate. 

Length | line. 

One half smaller than A. proximo, the thorax much less transverse, 
finely granulate, and the punctuation much more distant; the elytra 
without any basal depression, very closely punctured ; the legs 
entirely black. A. nigrita, Motsch., is described as "fere glabra" 
with pale legs. A. obscurata, Motsch., is much larger, also described 
as glabrous with testaceous tibiae and tarsi. In the present insect 
the two first joints of the antennae as well as the terminal ones are 
black, the intermediate joints more or less flavous. 

Dikoya. 

Aphthona dorsalis, Motsch. 

The description of the author, "Glabra, rufa, capite, thorace elytris- 
que dorso nigris, corpore subtus subinfuscata," agrees very nearly 
with two specimens before me. The antennae (the terminal joint 
excepted) and the legs are, however, testaceous, and the posterior 
femora have their posterior portion black ; this is not mentioned by 
Motschulsky. There are also very fine punctures visible at the elytra, 
when examined under a strong lens. It is therefore doubtful whether 
I am rightly referring these specimens to the present species. 

Obtained at the Horton Plains. 



1 887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 87 

EUCYCLA CEYLONENSIS, Sp. nOV. 

Fulvous ; antennae black, the basal joint fulvous ; thorax closely 
punctured, with or without a black basal spot; elytra strongly 
punctate-striate, the interstices finely punctuieJ, black, the suture 
narrowly and the apices more broadly fulvous. 

Var. a. Black, the first five joints of the antennae flavous. 

Var. b. Smaller ; piceous, the elytra fulvous. 

Length 1-lg line. 

Head nearly impunctate, fulvous. Antennae with the first joint 
long and slender, curved, the second short and thick, the four 
following joints still shorter and nearly equal in length, the rest 
widened and compressed. Thorax more than twice as broad as long, 
the posterior margin broadly produced at the middle, the sides 
straight ; the surface distinctly and very closely punctured, fulvous, 
sometimes with a central black spot widened at its base. Scutellum 
obscure fulvous. Elytra very convex, black with a greenish tint, 
strongly and regularly punctate-striate, the interstices very finely and 
sparingly punctured, the apices, in shape of a triangular spot, and 
the suture very narrowly and rather obscurely fulvous ; anterior 
coxal cavities open. 

The slender and elongate first joint of the antennae, the lobed 
thorax, punctate-striate elytra, and general rounded and convex shape 
seem to me to place the present insect in Baly's genus Eueycla. 
Thrijlaa of this paper has the general appearance and the punctate- 
striate elytra of the present insect, but may be known by the 
less transverse thorax and the much shorter basal joint of the 
antennae. 

LONGITARSUS LONGICORNIS, Sp. UOV. 

Ovate, convex ; obscure testaceous ; antennae much longer than the 
body ; thorax impunctate ; elytra scarcely visibly punctured ; the 
apices of the posterior tibiae piceous. 

Length 1 line. 

Head rather broader than long, entirely impunctate ; the frontal 
tubercles scarcely indicated ; labrum and palpi piceous. Antennae 
one half longer than the body, entirely testaceous, the third joint 
distinctly shorter than the fourth. Thorax about one half broader than 
long, the sides slightly rounded and constricted near the base, the 
anterior angles obliquely truncate, the surface entirely impunctate. 
Scutellum broader than long. Elytra ovate, narrowed near the base 
and the apices, extremely minutely punctured, only visible under a 
very strong lens, of a darker colour than the thorax ; underside 
and legs testaceous; the apical half of the posterior femora piceous ; the 
first joint of the posterior tibiae rather longer than the three following 
joints together. 

Bogawantalawa. 

The very long antennae, ovate shape, and the nearly impunctate 
upper surface are the distinguishing characters of L. longicornis. In 
one specimen the legs are of an entirely testaceous colour, but all the 
other characters are the same as in the type. 



88 MR. M. JACOB\' ON THE [Feb. 1, 

Parlina fulva, sp. nov. 

Oblong-ovate, fulvous ; apical joints of the antennae, the tibise, and 
tarsi piceous ; tliorax impunctate ; elytra closely punctured. 

Var. Entirely fulvous. 

Length 2 lines. 

Head impunctate, the frontal tubercles small but distinct, the 
carina short ; the penultimate joint of the palpi thickened, the apical 
one short, acutely pointed. Antennfe nearly as long as the body, the 
second joint short, the following joints nearly equal in length, the 
four basal ones fulvous, the rest more or less piceous. Thorax 
transverse, three times broader than long, the sides rounded and 
narrowly margined, the angles rather blunt, scarcely prominent, the 
surface with a transverse distinct groove near the base, not extending 
to the sides, scarcely visibly punctured or entirely impunctate. 
Scutellum triangular. Elytra without basal depression, closely and 
finely but distinctly punctured. The posterior tibiae mucronate ; the 
first joint of the posterior tarsi as long as the two following joints 
together ; claws appendiculate. Prosternum narrow ; the anterior 
coxal cavities open. 

Parlina was established by Motschulsky for the reception of a 
species of Ealtiea having the general characters of the genus Lactica, 
with which it agrees in the open coxal cavities and thoracic groove. 
The typical form (P. trancisa), which was also obtained by Mr. Lewis, 
differs from the species described here in its more general oval 
shape and more transverse thorax ; it agrees well enough with the 
description of the author to leave little doubt about the identity of 
the species. In Parlina the thoracic groove is placed close to the 
posterior margins and is bounded at the sides by a short, not very 
distinct perpendicular groove. P. fulva, of which several specimens 
are before me, differs in the almost entirely fulvous colour, in the 
much less transverse thorax, and in the narrower prosternum as well 
as in the move oblong shape. Chapuis seems to have overlooked the 
present genus, as he makes no mention of it in his ' Genera des 
Coleopteres.' 

From Lactica the type of Parlina differs in the much more trans- 
verse thorax and its sinuate groove, the latter not being bounded by 
a lateral depression. 

HyPNOPHILA VIOLACEIPENNIS, Sp. nOV. 

Ovate, very convex ; black ; basal joints of the antennae and the 
posterior tibiae obscure testaceous ; tliorax scarcely visibly punctured ; 
elytra dark violaceous, punctate-striate. 

Length | line. 

Head impunctate; the frontal tubercles obsolete. Antennae with 
the last five joints transversely dilated, black, the five lower joints 
testaceous. Ihorax transverse, three times as broad as long, the sides 
straight, the base with a very short longitudinal groove at each side ; 
the surface with a few very minute punctures, visible only under a 
strong lens. Scutellum piceous. Elytra very strongly convex, narrowed 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 89 

and rather pointed at the apices, each elytron with nine or ten rows 
of distinct punctures. Posterior tibijB dilated at the apices, the 
latter furnished witli a row of bristles and below with a long fulvous 
spine ; claws simple. 
A single specimen. 

Hypnophila apicipennis, sp. nov. 

Black ; base of the antennae, the head, thorax, and legs rufous ; 
eh'tra strongly puuctate-striate, the apices fulvous. 

Length ^ line. 

Head impunctate ; antennae with the last five joints transversely 
dilated, black, the others fulvous. Thorax nearly three times 
broader than long, transversely convex, rufous, shining, impunctate, 
the basal margin with a short longitudinal groove at each side. 
Elytra subglobose, very convex, strongly punctate-striate, their 
apices fulvous, this colour extending also partly to the sides. Claws 
simple. 

The single specimen obtained, like the following, is glued to a card, 
so that I cannot say anything about the underside. It is smaller 
than H. violaceipennis, but seems to possess all the characters of 
Hypnophila. 

Hypnophila rugicollis, sp. nov. 

Black ; head and thorax very finely rugose and wrinkled ; elytra 
dark purplish, distinctly punctate-striate. 

Length ^ line. 

Head finely rugose ; antennje black, of the same structure as in 
the preceding species. Thorax more than twice as broad as long, 
the sides defiexed ; the surface entirely covered with fine longitudinal 
rugosities, giving it an opaque appearance ; a small longitudinal 
indentation is placed on each side at the basal margin, and a lateral 
groove extends close to the lateral margin, the latter appearing 
somewhat thickened and shining. Elytra ovate, very convex and 
pointed at the apices, the punctures regular and moderately deep and 
placed in striae, the single punctures being very closely approached. 
Legs black, the posterior femora very strongly incrassate, their tibiae 
straight and armed at the apex with a long and distinct spine ; the 
first joint of the posterior tarsi nearly as long as the three following 
joints together ; claws simple. 

A single specimen. 

Manobia apicicornis, sp. nov, 

Piceous or black ; head, thorax, and legs fulvous ; antennae black, 
the four lower and the last joint fulvous ; elytra black, their apices 
fulvous, strongly punctate-striate. 

Var, a. Entirely fulvous. 

Var. b. Obscure piceous ; the tibiae fulvous. 

Length 1 line. 

Head impunctate, the frontal tubercles strongly raised, of an 
elongate triangular shape, bounded behind by a deep transverse 



90 MR. M. JACOBY ON THR [Feb. 1, 

groove, which extends to the inner margin of the eyes. Antennae 
nearly as long as the body ; the 4 or 5 lower joints obscure fulvous, 
the five following ones black, the apical joint reddish fulvous ; the 
third and fourth joints equal. Thorax transversely subquadrate, the 
sides straight, the posterior margin slightly lobed, the anterior angles 
obliquely truncate and slightly thickened ; surface with a deep, 
strongly sinuate, transverse groove near the base, extending nearly 
to the posterior angles, the latter produced into a tooth ; the disk 
impunctate, the groove itself with some punctures ; scutellum 
fulvous. Elytra with a well-marked basilar depression, the shoulders 
prominent, the disk strongly punctate-striate, the punctuation 
diminishing towards the apices, the interstices slightly costate near 
the sides. Prosternum rather broad ; the anterior coxal cavities 
open. 

31. apicicornis resembles greatly several species from the Malayan 
regions which served me for the establishment of the present genus ; 
their general appearance is that of a small species of Crepidodera, 
from which the open coxal cavities and the strongly sinuate thoracic 
groove separates Manohia. In M. apicicornis, which seems to be 
a very variable species, the apices of the elytra are pale fulvous, while 
the last joint of the antennae is of a more reddish colour ; this is 
constant in all the specimens before me, and separates the species 
from its allies. 

Crepidodera hirtipennis, sp. nov. 

Oblong-ovate ; black ; antennae and tarsi flavous ; thorax rugose- 
punctate ; elytra strongly punctate-striate, the interstices costate, 
and clothed with long white pubescence. 

Length | line. 

Head not visibly punctured, with some single long white hairs. 
Antennae a little shorter than the body, the third and the two follow- 
ing joints equal, slightly shorter than the second but not so stout ; 
terminal joints slightly thickened, the apical one fuscous, the rest fla- 
vous. Thorax rather more than twice as broad as long, the sides nearly 
straight, the disk strongly rugose-punctate, transversely grooved 
near the base. Elytra with regular rows of deep punctures, the 
interstices strongly costate, and furnished with long white single 
hairs. Legs black, tarsi flavous. 

Of this small and interesting little species there is only a single 
example before me. As the specimen is carded, I am not able to say 
anything about the state of the cavities or other characters of the 
underside, and have placed it at present in Crepidodera on account 
of the thoracic groove and punctate-striate elytra. The following 
species, of which also only a single specimen was obtained, is still 
smaller. Both agree in the almost equally stout femora of all the 
legs, which leaves it doubtful whether these species would not equally 
well find their place amongst the Galerucinse. 

Crepidodera minuta, sp, nov. 

Entirely pale fulvous ; head impunctate ; thorax very closely 



188/.] PHYTOPHAGOUS COLEOPTER.V OF CEYLON. 9l 

punctured ; elytra regularly punctate-striate, the interstices scarcely 
raised. 

Length h line. 

Rather smaller and narrower than the preceding species ; the 
thorax twice as hroad as long, the sides slightly narrowed towards 
the base, nearly straight, the surface extremely closely and dis- 
tinctly punctured, the basal groove distinct and placed close to the 
posterior margin ; elytra rather paler than the thorax, the punctures 
larger and arranged in regular rows ; all the femora thickened, the 
posterior ones scarcely more incrassate than the others. 

Sebaethe suturalis, sp. nov. 

Testaceous, terminal joints of the antennae fuscous ; head and 
thorax impunctate ; elytra very finely and closely punctured, a more 
or less distinct sutural stripe, narrowed behind, piceous. 

Length l|-2 lines. 

Head not longer than broad, impunctate ; the frontal tubercles 
strongly raised, transverse and nearly contiguous ; carina short but 
distinct. Antennae closely approached, two thirds the length of the 
body, the first three joints pale testaceous, the rest fuscous, the third 
joint one half longer than the second, but slightly shorter than 
the fourth joint. Thorax narrow, three times as broad as long, the 
sides slightly rounded and narrowly margined, the anterior angles 
thickened, the surface somewhat convex and entirely impunctate. 
ScutcUum rather broad, impunctate. Elytra slightly widened towards 
the middle, their apices rounded, the sides with a narrow margin ; 
the disk very finely and moderately closely punctured, obscure 
testaceous like the rest of the body, with a narrow posteriorly con- 
stricted sutural piceous stripe not extending to the apices ; the 
posterior tibiae longitudinally channelled; the first joint of the pos- 
terior tarsi as long as the two following joints together. 

Dikoya. 

Smaller than S. pallida, Jac, the sides of the thorax less strongly 
rounded, and the surface without depressions ; further distinguished 
by the sutural stripe ; this latter is, however, in some specimens 
scarcely visible, in others strongly marked. The colour of the 
antennae and that of the tarsi is also subject to variation, being 
sometimes obscure piceous and in some instances testaceous ; the 
impunctate thorax, the coloration of the elytra, in connection with 
the size, will help to separate S. suturalis from its allies. 

Sebaethe ceylonensis, sp. nov. 

Oblong-ovate, obscure testaceous ; thorax impunctate, the sides 
strongly rounded ; elytra extremely closely and finely punctured. 

Length 2|-3 lines. 

Head impunctate ; the eyes very large, divided by a space not 
broader than their diameter ; the frontal tubercles broadly trigonate, 
bounded behind by a deep groove. Antennae two thirds the length 
of the body, the joints slender and elongate, with the exception of 
the second, of nearly equal length. Thorax nearly three times as 



92 MR. M. JACOBY ON THE [Feb. I, 

broad as long, the sides rounded and nairowly margined. Elytra 
widened towards the middle, very closely and finely punctured ; the 
presternum narrow but distinct. 

Bogawautalawa. 

S. ceylonensis entirely resembles in regard to colour the uni- 
colorous variety of S. suturalis, but differs in being of much larger 
size and in having the sides of the thorax much more rounded ; the 
antennae have their joints also much more elongate, and the punc- 
tuation of the elytra is finer and more closely placed. As the four 
specimens before me all agree in the above characters, I must consider 
the species specifically distinct from the preceding. S. pallida, Jac, 
from Celebes, is another very closely allied species, but differs in the 
depressions of the thorax, the fulvous labrum, and the shorter 
antennae. 

SpHjERODERMA ORIENTALIS, Sp. HOV. 

Piceous ; the three basal joints of the antennae fulvous ; above 
reddish fulvous ; thorax very finely punctured ; elytra closely and 
finely semipunctate-striate. 

Length 1 line. 

Head impunctate ; the frontal tubercles distinct ; lower part of 
the face prominent, the anterior edge of the clypeus straight. 
Antennae about half the length of the body, black, the three lower 
joints fulvous, the second thinner and rather smaller than the 
preceding, the terminal jdints gradually thickened. Thorax trans- 
verse, three times broader than long, the sides straight, the posterior 
margin distinctly sinuate at each side, the median lobe slightly 
produced and rounded, the surface finely and evenly punctured. 
Scutellum small. Elytra very convex and distinctly narrowed 
towards the apices, the shoulders not prominent, the surface very 
closely and somewhat more distinctly punctured than the thorax, 
the punctuation arranged in semiregular rows. Legs [)iceous. Pro- 
sternum longer than broad. Elytral epipleurae broad, nearly extend- 
ing to the apices. 

Galle. 

I am unable to say whether the present species is identical with 
one or the other described by Motschulsky. In some specinieus the 
thorax is more or less stained with piceous. 

Chabria (gen. nov. Halticinorum). 

Anterior coxal cavities open. Body ovate, roundi d, very convex. 
Antennae widely separated, filiform, shghtly thickened towards the 
apical joints. Thorax narrowly transverse, four times as broad as 
long, the sides rounded. Scutellum triangular. Elytra irregularly 
punctured, convex, strongly deflected towards the apices, their epi- 
pleurse broad and continued below the middle. Posterior femora 
strongly incrassate ; tibiae not channelled, the posterior ones with a 
distinct spine ; the first joint of the posterior tarsi as long as the 
two following joints together ; claws appendiculate. Presternum 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 93 

narrow but distinct, longer than broad, its base slightly widened and 
rounded. 

The general apjiearance of the insect for which I am obliged to 
propose the present genus has much the appearance of a species of 
Chrysomela on account of the ovate and strongly convex shape and 
the very transversely shaped thorax : the s[)ace between the insertion 
of the antenniE is much greater than is the case in the majority of 
the Halticince, while the strongly incrassate posterior femora leaves 
no doubt as to the proper place of the genus amongst the latter 
family. 

Chabria nigroplagiata, sp. nov. (Plate X. fig. 8.) 

Black ; antennjfc and the tibise flavous ; above fulvous or flavous, 
a spot at the vertex, two at the thorax, two transverse bands at the 
elytra, and a triangular spot near the apices of the latter black. 

Length 2^ lines. 

Head impuuctate, the vertex with a black spot (sometimes wanting); 
palpi piceous as well as the apices of the jaws ; antennae entirely 
flavous or fulvous, scarcely half the length of the body, the third 
joint about one half longer than the second, the terminal joints 
gradually and slightly thickened. Thorax more than three times 
broader than long, tlie sides rounded, the anterior angles obliquely 
truncate (in one specimen more pointed), the surface with a few 
scarcely visible punctures, fulvous, with a black spot of variable 
shape placed on each side at the base. Scutellum black, or with the 
base only of that colour. Elytra very convex, rounded towards the 
middle and pointed at the apices, their surface scarcely or not visibly 
punctured ; each elytron with a broad concave-shaped band near the 
base, not quite extending to either margin ; another transverse band is 
placed at the middle, each end being widened, and a triangular-shaped 
spot near the apex. Underside and the femora and tarsi black. 

Bogawantalawa. 

Chabria apicicornis, sp. nov. (Plate X. fig. 9.) 

Piceous ; antennae testaceous, the two apical joints black ; thorax 
and elytra minutely punctured ; the posterior tibiae and the tarsi 
testaceous or flavous. 

Var. The basal joints of the antennae piceous. 

Length 2^-3 lines. 

Head rather broader than long, impunctate, the frontal tubercles 
almost entirely absent, the space between the antennae broad, divided 
at the base by a rather deep groove and bounded behind by another 
transverse groove; clypeus broad, scarcely narrowed above, and 
forming a single piece with the face ; labrum transverse, with a 
row of fine punctures ; palpi incrassate at the penultimate joint, 
the apical one acute and conical. Antennae half the length of the 
body, the third and fourth joints equal, the following slightly widened 
at their apices, the two or three terminal joints black, the others 
testaceous. Thorax with the sides very strongly rounded and narrowly 
margined, the anterior angles entirely obsolete and oblique and 



94 MR. M. JACOBY ON THE [Feb. 1, 

indicated only near the middle by a thickened rounded fovea ; the 
surface scarcely visibly punctured. Elytra very convex, deflexed and 
pointed at the apices, without any basal depression or prominent 
shoulders, the surface punctured like that of the thorax. Body 
below coloured like the upper surface ; the tibiae and the tarsi more 
or less distinctly flavous. 
Dikoya, Bogawantalawa. 

Ph.elota (gen. nov. Halticinorum). 

Body ovate, convex, subcylindrlcal, pointed posteriorly. Antennae 
rather distant, gradually thickened at the terminal joints. Thorax 
transverse, the sides roujided, the anterior angles thickened and 
obliquely rounded, the disk without depression. Elytra punctate- 
striate. Anterior tibife unarmed ; the posterior ones with a very 
short spine, somewhat widened at their apices, and with a short and 
shallow longitudinal depression or groove ; the first joint of the 
posterior tarsi as long as the two following joints together ; claws 
appendiculate. Presternum broad, its base truncate. Mesoster- 
num of the same shape but half the size. Anterior coxal cavities 
closed. 

Phalota, like Gkabria, has much the general appearance of a spe- 
cies of Chrysomela ; from Chabria it differs in the shorter antennae, 
the punctate-striate elytra, and the closed coxal cavities. 

PHiEI-OTA SEMIFASCIATA, Sp. UOV. 

Fulvous or piceous ; above obscure greenish aeneous varied with 
fulvous ; thorax minutely punctured, the disk greenish ; elytra 
regularly punctate-striate, greenish aeneous, the interstices alternately 
more or less obscure fulvous. 

Far. Above and the legs entirely fulvous (immature ?). 

Length 2 lines. 

Head with a few minute punctures ; eyes large ; frontal tubercles 
transversely trigonate, nearly contiguous ; the carina indistinct ; 
labrum more or less fulvous. Antennae half the length of the body, 
the five lower joints fulvous, the rest black, the apex of the terminal 
joint fulvous ; the second to the fifth joints short, nearly equal, the 
six terminal ones thickened, shghtly longer than broad. Thorax 
transverse, the anterior margin nearly straight, the posterior one and 
the sides slightly rounded, the anterior angles much thickened and 
obtusely rounded, the surface minutely punctured, greenish aeneous, 
the marginsmore or less fulvous. Elytra very convex, subcyUndrical, 
and pointed at the apices, strongly punctate-striate. Legs piceous 
or fulvous, more or less stained with greenish aeneous. 

Bogawantalawa. 

In some specimens the elytra show alternate longitudinal bands of 
aeneous and fulvous. 

Pexodorus (gen. nov. Halticinorum). 
Bodv ovate, widened behind ; palpi slender, filiform. Antennae 
filiform^ the second joint short, the third and fourth joints equal. 



J837.] PHYTOPHAGOUS COLEOPfERA OF CEYLON. 95 

Thorax narrowly transverse, three times as broad as long. Scutellum 
triangular. Elytra with a more or less distinct basal depression, 
semipunctate-striate, their epipleurtB not continued below the middle ; 
the four anterior tibise unarmed, the posterior ones with a small 
spine ; the first joint of the posterior tarsi as long as the two 
following ones together; claws appendiculate. Prosternum distinct; 
the anterior coxa! cavities closed. 

Pexodorus will enter Chapuis's eighth group, the Oxygonince, on 
account of the closed coxal cavities and the narrow transverse 
thorax. From Oxygona the genus is distinguished by the short 
ovate general shape of its body and the elytral epipleurse, which are 
obsolete below the middle. 

Pexodorus ceylonensis, sp. nov, (Plate X. fig. 10.) 

Black or metallic green ; the basal joints of the antenna?, the 
four anterior legs, and the posterior tibioe testaceous ; thorax 
impunctate ; elytra finely and closely punctate-striate. 

$ (?). Elytra with deep basal depression ; antennae entirely 
testaceous ; all the femora and the posterior tibiae black. 

Length l|-2 lines. 

Head rather broader than long ; the frontal elevations ovate, but 
little raised and small ; eyes entire, of oblong shape ; clypeus rather 
flat ; labrum obscure fulvous. Antennae two thirds the length of the 
body, testaceous, the three or four apical joints fuscous. Thorax 
rather more than three times as broad as long, the posterior margin 
slightly rounded, the sides narrowly margined, nearly straight, the 
anterior angles somewhat broad and slightly produced, furnished as 
well as the posterior ones with a single hair ; the surface entirely 
impunctate. Elytra widened towards the apices, with a shallow 
basal depression ; the surface finely and very closely punctate-striate; 
the posterior femora strongly incrassate, piceous, the others and the 
tibiae and tarsi testaceous. Prosternum narrow ; mesosternura 
much broader, its base slightly emarginate. 

Dikoya, 

I am not quite sure whether I rightly refer two specimens, which 
differ from the others in the very deep elytral depressions and 
diiferently coloured legs, to the female sex. lu the absence of other 
distinguishing characters it is as well to regard them as such. Some 
specimens are of a metallic greenish or bluish colour and the antennae 
are entirely flavous; in others the colour of the latter is nearly black; 
but I cannot discover any other differences of importance. 

Philogeus (gen. nov. Halticinorum). 

Ovate, convex, subcylindrical. Anterior coxal cavities open ; eyes 
large ; palpi robust ; antennae with dilated apical joints. Thorax 
transversely subquadrate, with a shallow transverse groove near the 
base, the anterior angles oblique, notched before the middle. 
Elytra finely punctate-striate, not depressed below the base. 
Posterior femora strongly incrassate, their tibiae dilated at the apices, 
with a shallow longitudinal groove, armed with a spine ; the first 



96 MR. M. JACOBY ON THE [Feb. 1, 

joint of the posterior tarsi rather longer than the two following 
joints together ; claws bifid ; prosternum longer than broad, very 
distinct ; mesosternum transversely subquadrate. 

The insect for which I am obliged to establish the present genus 
has quite the appeaiance of a small species of Typophorus amongst 
the Eumolpiiise. From any other genus of the present family, 
especially from Munobia, Jac, Philogeus may be distinguished by the 
dilated and flattened posterior tibiae in connection with the thoracic 
groove and tiie dilated antennae, as well as by the bifid claws, 
which is a character of rare occurrence amongst the Halticinse. 

Philogeus fulvipennis, sp. nov. 

Fulvous ; head, antennae, thorax and the legs black ; thorax 
impunctate ; elytra very finely puactate-striate, fulvous. 

Length \\ line. 

Head impunctate, the frontal tubercles obsolete ; the carina 
acutely raised ; antennae closely approached, nearly as long as the 
body, the third and fourth joints equal, slightly longer than the 
second, which is thickened, the sixth to the tenth joints gradu- 
ally and distinctly widened, pubescent, the terminal one of usual 
size, often fulvous. Thorax about one half broader than long, the 
sides straight, the posterior margin slightly and broadly rounded, 
the anterior angles forming an obtuse tooth before the middle ; the 
surface scarcely visibly punctured, with an obsolete sinuate transverse 
groove near the base extending some way upwards at the sides ; 
scutellum rather broad, its apex rounded. Elytra extremely finely 
and rather distantly punctate-striate, their apices rounded. 

Dikoya. 

Amphimeloides (gen. nov. Halticinorum). 

Subovate, convex. Antennae separated, inserted immediately 
below the eyes, short, their apical joints widened. Thorax trans- 
verse, the sides angulate before the middle. Elytra irregularly 
punctured. Posterior femora strongly incrassate, their tibiae dilated 
and slightly longitudinally sulcate near the apices, the latter armed 
with a long spine. Claws appendiculate. Prosternum very narrow 
but distinct. Anterior coxiil cavities open. 

From all other genera of Halticidae with the exception of Atnphi- 
mela the present genus differs by the broad space dividing the 
insertion of the antennae, while the open coxal cavities will distin- 
guish the genus from Amphimela proper. 

Amphimeloides dorsalis, sp. nov. 

Fulvous ; the apical joints of the antennae, the posterior femora, 
and the breast piceous or black ; thorax scarcely visibly, elytra 
more distinctly and closely punctured, each elytron with a broad 
longitudinal black band, abbreviated posteriorly. 
■ Length 1 line. 

Head impunctate, without transverse groove or frontal elevations ; 
the cly pens not separated from the face, which forms a plane surface; 



1887.] PHYTOPHAGOUS COLEOPTERA. OF CEYLON. 97 

labrum piceous ; palpi long aud slender. Antennse inserted close to 
the inner margin of the eyes, scarcely extending in length to the 
base of the thorax, the second joint short and thickened, the third 
more slender and longer, the rest gradually widened and trans- 
versely shaped ; black, the four basal joints fulvous. Thorax at 
least three times broader than long, pale fulvous, the sides straight 
and forming a distinct angle before the middle, the posterior margin 
evenly and moderately rounded ; the surface without depressions, 
smooth and nearly impunctate. Scutellum broadly ovate, black. 
Elytra convex, subcylindrical, closely and distinctly punctured, 
fulvous, each elytron with a broad black band commencing at the 
middle of the base and extending below the middle, the outer 
margin deeply concave at the middle. Legs fulvous, the posterior 
femora piceous as well as the sides of the breast. 

The elytral band is slightly widened at the apex, and approaches 
gradually towards the suture without, however, touching tlie latter. 

Tegyrius (gen. nov. Halticinorum). 

Ovate, subcylindrical. Antennae slender, filiform, the third joint 
slightly longer than the second. Thorax transversely subquadrate, 
the surface transversely but obsoletely grooved near the base. 
Elytra convex, broader than the thorax, without depressions, finely 
and semiregularly punctured. Posterior femora strongly incrassate, 
their tibiae dilated and longitudinally channelled, their apices with a 
small spine ; the first joint of the posterior tarsi as long as the three 
following joints together. Claws appendiculate. Prosternum 
broad, subquadrate. Mesosternum broader than long, its base 
concave-emarginate. Anterior coxal cavities open. 

Tegyrius has the general shape and appearance of Philogeus, but 
differs in the filiform antennae, the much longer metatarsus of the 
posterior tarsi, and in the appendiculate, not bifid, claws. From 
Longitarsus the genus may be distinguished by the transverse 
sinuate groove of the thorax and the broad prosternum. 

Tegyrius metallicus, sp. nov. 

Black ; antennae, legs, the posterior femora excepted, testaceous ; 
above metallic greenish aeneous ; head and thorax impunctate ; 
elytra very finely semipunctate- striate. 

Length 1 line. 

Head impunctate ; the frontal tubercles and the carina very 
narrow and rather indistinct. Antennae nearly as long as the body, 
the fourth joint one half longer than the third, the second thickened. 
Thorax scarcely twice as broad as long, the sides straight, the 
anterior angles obliquely truncate and shghtly thickened, the basilar 
transverse groove sinuate, not very deep and not extending to the 
sides ; the disk entirely impunctate. Elytra convex, subcylindrical, 
without basal depression, the shoulders not prominent, the apices 
rounded ; surface very closely and minutely punctured, the punctua- 
tion arranged in semiregular rows ; the anterior tarsi, the posterior 
femora, and the inner side of the posterior tibiae blackish. 

Proc. Zool. Soc— 1887, No. VTL 7 



98 Ma. M. JACOBY ON THE [Feb. 1, 

The antenn?e and the legs in this r-pecies are subject to variation 
in colour ; sometimes the first two joints of the antennae are stained 
with piceous as well as the legs and tarsi, the latter in other speci- 
mens being entirely testaceous with the exception of the posterior 
femora. 

Alytus (gen. nov. Halticinorum) . 

Body ovate, pointed behind. Head not longer than broad ; fronta 
tubercles strongly raised ; antennae as long as or longer than the body, 
the second and third joints nearly equal. Thorax subquadrate, the 
angles not produced, the surface with a distinct transverse groove 
near the base, extending to the posterior angles. Scutellum trigonate. 
Elytra ovate convex, pointed posteriorly, regularly punctate-striate. 
Posterior femora strongly incrassate, their tibiae with a distinct spine ; 
the first joint of the posterior tarsi as long as the two following 
joints together ; claws appendiculate. Anterior coxal cavities open. 
Piosternum narrowly elongate, much longer than broad ; meso- 
sternum distinct, subquadrate. 

In its general appearance the insect, for the reception of which I 
am obliged to establish the present genus, resembles a species of 
Longitarsus, from which the distinct transverse groove of the thorax 
and the short metatarsus of the posterior legs separate it ; the 
punctate-striate and the ovate and strongly pointed elytra are further 
characteristic of Alytus. A single species is before me. 

Alytus ceylonensis, sp. nov. 

Fulvous or testaceous ; posterior femora piceous ; the disk of the 
thorax impunctate, the groove punctured ; elytra strongly punctate- 
striate. 

Length |-1 line. 

Head impunctate, the frontal tubercles strongly developed, elon- 
gate ; the third joint of the antennae thinner than the second, but 
scarcely longer, the following joints more slender and elongate. 
Thorax transversely quadrate, slightly constricted at the base, the 
sides nearly straight, the angles rather obtuse, the surface entirely 
impunctate, the transverse groove closely punctured. Elytra ovate, 
convex, widened at the middle, fulvous ; each elytron with about ten 
rows of regular and distinct punctures, the first row very short. Legs 
testaceous, the apices of the posterior femora piceous. 

The general colour of the upper surface is dark fulvous, very 
shining, the thorax and the legs being of a paler tint. The elytra 
are strongly narrowed at the base and at the apices, so that the 
thorax is broader than the elytra at the base. 

Thryl^a (gen. nov. Halticinorum). 

Body rounded, subovate ; eyes rather large ; frontal tubercles in 
shape of oblique narrow ridges. Antennae rather short, the ter- 
minal joints thickened. Thorax transverse, the anterior angles 
obliquely truncate, the surface without transverse groove, the basal 



99 



J 887.] PEIYTOPHAGOUS COLEOPTERA OF CEYLON. 

margin with a short perpendicular groove. Elytra distinctly 
punctate-stnate ; their epipleurae very broad, concavei and continued 
to the apices; posterior femora strongly thickened; tibije not 
channelled, the posterior ones with a short spine ; the first joint of 
the posterior tarsi as long as the two following joints together ; 
claws appendiculate. Prosternum broad, one half longer than broad. 
Me^osternum transversely quadrate. Anterior coxal cavities open. 

Ihe rounded convex shape, the short perpendicular basal grooves 
ot the thorax, in connection with the punctate-striate elytra, the 
broad prosternum, and open coxal cavities separate this genus. 

Thryl^a variabilis, sp. nov. 

Reddish fulvous; head, thorax, and the legs black; thorax 
distinctly punctured ; elytra strongly punctate-striate, dark fulvous. 
yar. J^ulvous; elytra black, the suture and the apices fulvous. 
Length 1 line. 

Head impunctate, the clypeus thickened ; labrum margined with 
fulvous. Antennae about half the length of the body, black, the 
apices of the first five joints stained with fulvous ; the' second ioint 
as thick, but half the size of the first, the third and the two follow- 
ing joints nearly equal, shorter and thinner, the rest more flattened 
and dilated. Thorax transverse, rather more than twice as broad as 
long, the sides straight, the anterior angles oblique forming a 
thickened angle before the middle; surface finely but distinctly and 
not very closely punctured, black. Scutellum fulvous. Elytra 
rounded, strongly and regularly punctate-striate, the interstices with 
a tevv fine punctures, slightly convex near the sides and apices, the 
shoulders thickened and somewhat prominent. Legs black : tarsi 
piceous. o , oi 

Bogawantalawa. 

MoRYLUs (gen. nov. Halticinorum). 

Body ovate Antennae slightly widened at the terminal joints, 
the third and fourth joints equal. Thorax transverse, without de- 
pressions, the anterior angles oblique. Scutellum broader than Ion- 
Elytra irregularly punctured, their epipleurae extending below the 
middle. The posterior femora strongly incrassate, their tibia, 
deeply longitudinally channelled, armed at the apices with a small 
spine; the anterior tibiae unarmed. Claws appendiculate. Pro- 
steruum very broad. Mesosternum more than twice as broad as 
long. Anterior coxal cavities open. 

Mor^lusJl^vees^^'ith>Sebae(ke, Baly, in the deeply sulcate posterior 
tibiae, but diiiers in the shape of the thorax, the sides of which are 
straight, not flattened or margined, and in the broad prosternum. 

MORYLUS FULVIPENNIS, Sp. nov. 

Black, head and thorax impunctate; elytra fulvous, depressed 
below the base, very closely and distinctly punctured ; legs black 
Length 1| line. ° 

Head entirely impunctate, the frontal elevations in shape of narrow 

7* 



100 MR. M. JACOBY ON THE [Feb. 1 , 

oblique ridges, wliich continue between the antennae. ; the lower part 
of the face concave ; the penultimate joint of the palpi transversely- 
quadrate. Antenna rather robust, more than half the length of the 
body, black ; the first joint thicic and rather short, the second as 
thick but one third shorter, the two following joints elongate and 
thinner, the rest gradually and moderately dilated, longer than 
broad. Thorax rather more than twice as broad as long, widened 
at the middle, the sides perfectly straight, the anterior angles obliquely 
cut, forming a tooth before the middle, the surface not visibly punc- 
tured. Elytra slightly widened towards the middle, broadly and 
rather obsoletely depressed below the base, the shoulders some- 
what prominent ; the surface very closely and distinctly punctured, 
reddish fulvous. 

IvALiA (gen. nov. Halticinorum). 

Body very convex, ovate. Antenn?e with the seven last joints 
transversely dilated. Thorax transverse, without grooves. Elytra 
irregularlv punctured ; the posterior femora strongly incrassate ; 
posterior tibiae stout, widened behind, dee[)ly channelled, the edges 
armed with several teeth, the apices with a long spine. Claws appen- 
diculate. Prosternum narrowly elongate ; niesosternum very trans- 
versely shaped, widened at the middle. Anterior coxal cavities open. 

I am obliged to propose the present genus for the reception of 
some small species of Halticidse having the general shape and 
appearance of Apteropeda, but differing from that and the allied 
genera placed by Chapuis in his 16th group by the irregularly 
punctured elytra and the appendicnlate claws. 

IVALIA VIRIDIPENNIS, sp. nov. (Plate X. fig. 12.) 

Reddish fulvous ; terminal joints of the antennae black ; elytra 
metallic green, finely and closely punctured, their apices very pointed. 

2 C?). Larger, more rounded, the apices of the elytra fulvous. 

Length |-1 line. 

Head impunctate, fulvous ; the frontal tubercles obsolete ; palpi 
long and rather slender, the apical joint piceous. Antennae black, 
the four lower joints testaceous, the third and fourth joints small, 
equal, the following transversely dilated, pubescent. Thorax nearly 
three times as broad as long, widened at the middle, the sides nearly 
straight, the angles obsolete; surface extremely finely punctured, 
fulvous. Elytra ovate, strongly narrowed and pointed at the apices, 
the surface very closely punctured. Legs fulvous ; the posterior 
tibise with three or four teeth at their margins. 

Bogawantalawa. 

\\\ the specimen which I consider to be the female of this species 
the general sha[)e is more robust, and the elytra have their extreme 
apices of a fulvous colour ; the teeth at the tibiae are not so plainly 
marked; but other differences of importance I cannot find. 

IVALIA METALLICA, Sp. UOV. (Plate X. fig. II.) 

Black or piceous below ; the four basal joints of the antennae 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 101 

testaceous ; thorax metallic blue or greenish, finely punctured ; 
elytra metallic purplish or cupreous, closely punctured. 

Length 1 line. 

Head impunctate ; the frontal tubercles feebly raised. Antennte 
black, the last seven joints transversely dilated, "the others obscure 
testaceous. Thorax transversely snbcylindrical, widened towards the 
middle, the surface finely granulate and punctured, with an obscure 
longitudinal impression near the lateral margin. Scutellum triausnlar, 
piceous. Elytra very convex, narrowed behind, of a reddish metallic 
cupreous colour, very closely and irregularly punctured. Legs black, 
the tibiae rather lighter, the posterior ones armed with three or 
four teeth ; their apices with a long fulvous spine ; claws appen- 
diculate. 

Bogawantalawa. 

This species is of the same shape as the preceding, with which it 
has further all the structural characters in common ; the different 
coloration sufficiently distinguishes it. 

IVALIA FULVIPENNIS, sp. HOV. 

Black ; head and thorax impunctate ; elytra dark fulvous, very 
closely and irregularly punctured. 

Length 1 line. 

Head broader than long. Antennae short, robust, the three lower 
jomts obscure fulvous, the rest black. Thorax three times as broad 
as long, the sides nearly straight, surface entirely impunctate, black. 
Scutellum black. Elytra rounded, very convex, dark reddish fulvous, 
closely punctured, the interstices somewhat rugose or wiinkled. Legs 
piceous, the tibiae more or less obscure fulvous. 

Demarchus (gen. nov. Halticinorum). 

Body ovate ; pubescent. Antennae filiform, the third joint more 
than double the length of the second ; palpi robust. Thorax trans- 
verse with an anterior and posterior transverse depression. Scutellum 
subpentagonal. Elytra pubescent, finely rugose, their epipleurae 
disappearing below the middle. Tibiae simple, unarmed ; the first 
joint of the posterior taisi not longer than the second ; claws ob- 
soletely bifid. Anterior coxal cavities open. Prosternum scarcely 
visible. iMesosternum narrow and pointed. 

The single specimen before me, upon which I am obliged to 
estabhsh the present genus, resembles in general appearance a species 
of the genus Sebaetha, from which the simple tibije, transversely 
impressed thorax, and the pubescent elytra will at once distinguish 
It. The posterior femora are moderately but very distinctly iu- 
crassate. 

Demarchus pubipennis, sp. nov. 

Testaceous ; head rugose ; thorax shining, nearly impunctate ; 
elytra obscure fulvous, the basal and the lateral margin obscure piceous.' 
Length 2 lines. 
Head finely rugose at the vertex, the frontal tuborclea distinct. 



102 MR. M. JACOBY ON THE [Feb. I, 

nearly square-shaped and smooth ; labrum piceous. Antennae nearly 
as loiig as the body, testaceous, the third and following joints elon- 
gate, nearly equal in length and much longer than the second joint. 
Thorax more than twice as broad as long, the sides rounded at the 
middle, narrowed near the anterior angles, the latter slightly pro- 
minent ; the surface shining, scarcely visibly punctured near the side, 
with a short anterior and posterior transverse groove and a small 
anterior fovea as well as an obsolete oblique posterior lateral depres- 
sion. Scutellum obscure fulvous. Elytra closely pubescent, very 
finely ruoose-punctate, the basal margin and the sides to below the 
middle obscure piceous or fuscous, the rest of the surface very ob- 
scure fulvous. Legs and underside testaceous ; the claws indistinctly 
bifid. 

SpHjEropleura (gen. nov. Halticinorum). 

Body strongly rounded and convex. Antennae filiform, the first 
joint slender, the second and following joints nearly equal and shorter. 
Thorax suhhemispherical, without dejjression. Elytra puuctate- 
striate, their epipleurfe very broad and concave, continued to below the 
middle ; the posterior femora strongly dilated ; the four posterior 
tibiae mucronate ; the first joint of the posterior tarsi as long as the 
two following joints united ; claws appendiculate. Prosternum nar- 
row, deeply longitudinally channelled. Mesosternum deeply emargi- 
nate at its apex. Anterior coxal cavities closed. 

Sphceropleitra agrees in shape, which resembles that of a species 
of Coccinella, with Sphcerophyma, Baly, Argopistes, Motsch., and 
Homelea, Jac. It differs from the first-named genus in the much 
less strongly dilated posterior femora, the longer and not dilated 
tibiae, and the smaller eyes, also in the want of a thoracic median 
lobe. From Argopistes this genus may be separated by the filiform 
antennae and the punctate-striate elytra ; and from Homelea by the 
much more strongly incrassate posterior femora, the latter in i/o?ne/e« 
being only about one half thicker than the rest of the thighs, and 
the mesosternum strongly transverse and of different shape, and the 
elytra irregularly punctured. 

Sph.eropleura tricostata, sp. nov. 

Piceous below ; antennas, tibia3, and tarsi testaceous ; head and 
thorax impunctate, black ; elytra finely punctate-striate. 

$ . Each elytron with three short but distinct costae near the apex. 

Var. Elytra fulvous. 

Length 1| line. 

Head entirely impunctate ; the frontal tubercles very small and 
indistinct; labrum fulvous. Antennae about half the length of the 
body, testaceous, the terminal joint fuscous. Thorax suhhemi- 
spherical, more than three times as broad as long, the sides straight, 
the angles not produced but distinct. Elytra wider at the base than 
the thorax, very convex and roimded, the punctuation very fine at 
the middle of the disk, much more distinct towards the sides ; of the 
three costae placed near the apex in the female, the first is shorter 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 103 

than the others, but none extend to the end of the elytra nor to the 
middle. The shoulders are but little prominent, and between them 
and the lateral margin of the elytra there is a broader impuuctate 
space extending to the middle, the lateral margin itself being accom- 
panied by a row of deep punctures. The legs are subject to some 
variation in colour, being sometimes dark fulvous, but the anterior 
tibiae seem to remain testaceous. 

AuLACOPnoRA sTEVENsi, Baly. (Plate XI. fig. I.) 

Testaceous ; head and thorax impunctate ; elytra finely punctured; 
a sutural and submargiual narrow stripe, as well as their extreme 
apices, black. 

d • The fourth joint of the autennse strongly swollen and elongate. 

Length 3 lines. 

Head rather swollen at the vertex, impunctate, the frontal tubercles 
narrowly transverse. Antennae more than two thirds the length of 
the elytra, entirely testaceous, the second joint extremely short, the 
third and fifth joints triangularly dilated in the male, the fourth 
greatly enlarged in the same sex, the rest of nearly equal length and 
slender. Thorax transverse, the sides rather deflexed and widened 
towards the apex, the surface with a deeply impressed transverse 
groove near the base, impunctate. Scutellum black. Elytra rather 
convex and gradually widened posteriorly, extremely finely and rather 
closely punctured, a narrow sutural stripe extending to the apices 
and round the latter, and another equally narrow stripe near the 
lateral margin, commencing at the base and alibreviated before the 
apex of each elytron, black. Legs and underside entirely testaceous. 

The female only differs from the male in having simple antennae. 

Bogawantalawa. 

AuLAcoPHORA NiGRiPETA, Duviv. (Plate XI. figs. 2, 3.) 

PhYLLOBROTICA HIRTIPENNIS, Sp. UOV. 

Obscure piceous or black below ; basal joints of the antennae, the 
head, thorax, and the femora fulvous ; elytra obscure testaceous, 
finely rugose and pubescent, the margins narrowly fuscous. 

Length Ig line. 

Head impunctate ; the frontal tubercles distinct but small ; palpi 
robust. Antennae two thirds the length of the body, piceous, the 
three basal joints fulvous. Thorax transverse, the sides slightly con- 
stricted at the base, the disk rather deeply transversely depressed, 
impunctate. Elytra very obscure pale or darker testaceous, sometimes 
fuscous, the suture and the lateral margin at the shoulders darker ; 
the surface slightly rugosely punctured and moderately closely covered 
with stiff whitish hairs ; their epipleurae extremely narrow. Tibiae 
unarmed, fuscous ; the first joint of the posterior tarsi as long as the 
two following joints together ; the anterior coxal cavities open. 

Nuwara Eliya. 

The legs are sometimes entirely fulvous, but generally the femora 
only are of that colour. 



104 MR. M. JACOBY ON THE [Feb. 1, 

Phyllobrotica marginata, sp. nov. 

Testaceous ; the basal joints of the antennae, the head, and thorax 
fulvous ; seutellum black ; elytra testaceous, very finely rugosely 
punctate, the sutural and lateral margins narrowly black. 

Length 2 lines. 

Head impunctate ; the frontal tubercles trigonate, strongly 
raised. Antennse nearly as long as the body, the third joint double 
the length of the second, the following ones nearly equal to tiie 
third joint, the three basal ones fulvous, the rest fuscous. Thorax 
transverse, more than twice as broad as long, the anterior and 
posterior margin straight and parallel ; the surface rather deeply 
transversely depressed, impunctate, shining, fulvous like the head. 
Seutellum black. Elytra parallel, extremely finely punctured and 
rugose, the extreme lateral and sutural margins black ; their epipleurse 
very narrow. Legs slender, the tibise unarmed ; the first joint of 
the posterior tibise as long as the two following joints together ; 
claws appendiculate ; anterior coxal cavities open. 

P. marginata differs somewhat from the more typical species of 
the genus in the more transversely shaped thorax and the distinct, 
although very narrow elytral epipleurse. In the absence of other 
marks of distinction, I have placed the species in Phyllobrotica. 

MiMASTRA CAPITATA, Sp. UOV. 

Narrowly elongate ; testaceous ; antennse and the breast black ; 
vertex of the head metallic greenish ; thorax impunctate, depressed 
at the disk ; elytra finely punctured and rugose, with a metallic 
green gloss, the margins narrowly greenish black. 

Length 2-3 lines. 

Head impunctate at the vertex, the latter very finely granulate, 
metallic green or seneous ; lower part of the face testaceous ; the 
labrum obscure piceous. Antennse longer than the body, the second 
joint extremely short, the others very elongate and of nearly equal 
length ; black, sometimes the basal joints obscure fulvous. Thorax 
square-shaped, the disk deeply transversely depressed, impunctate, 
testaceous or fulvous, the lateral margins more or less stained with 
dark greenish. Elytra finely transversely rugose and closely punc- 
tured ; testaceous, stained with metallic greenish, the latter colour 
more distinct at the basal margin ; the suture and the lateral 
margins also narrowly dark seneous, which colour extends also to 
the epipleurse ; the underside either entirely or partially black. 
Legs fulvous, the tarsi generally fuscous ; the first joint of tlie 
posterior tarsi as long as the three following joints together ; tibise 
unarmed. Anterior coxal cavities open. 

Dikoya. 

Except in the greater length of the posterior metatarsus, this 
species does not diifer in any other material way from its allies. 

MiMASTRA ROBUSTA, Sp. UOT. 

Below piceous or blackish ; above fulvous ; antennse and legs 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 105 

flavous ; thorax impunctate, obsoletely depressed ; elytra very 
minutely punctured. 

Length 2| lines. 

Head impunctate, not longer than broad ; the frontal tubercles in 
shape of narrow transverse ridges ; the anterior margin of the clypeus 
straight ; extreme apices of the jaws black. Antennae slender, nearly 
as long as the body, flavous, the two apical joints slightly fuscous, 
the third joint slightly shorter than the fourth. Thorax subquadrate, 
scarcely more than one half broader than long, the sides narrowed 
towards the base, the angles not prominent, the surface feebly im- 
pressed at the middle, impunctate. Elytra rather flattened, the 
base scarcely or feebly raised, the surface very finely and rather 
closely punctured, the punctuation here and there arranged in rows ; 
elytral epipleurae continued below the middle. Legs rather short 
and stout, flavous ; tibiae unarmed; the first joint of the posterior 
tarsi as long as the two following joints together ; claws appendicu- 
late. Anterior coxal cavities incomplete. Underside blackish, with a 
slightly metallic bluish gloss. 

Although the general shape of this species is less elongate and 
the thorax less transversely shaped than is the case with most of 
the other representatives of the genus Mimastra, it would scarcely be 
deemed sufficient to estabhsh on these differences alone another 
genus, as, moreover, the open coxal cavities and the unarmed tibise 
are characteristic of the present genus. 

Galerucella virida, sp. nov. 

Oblong, rather convex; testaceous; antennae, tibiee, and tarsi 
fuscous ; above pale green, closely punctured ; the head, thorax, and 
the elytra narrowly margined with flavous. 

Length 4 lines. 

Head very finely and closely punctured, with a fine longitudinal 
central groove, the vertex green, the sides narrowly flavous ; the 
anterior margin of the clypeus straight ; labrum and palpi piceous. 
Antennae inserted just above the lower edge of the clypeus, about 
half the length of the body, fuscous, the third joint one half longer 
than the second, the following ones slightly longer. Thorax three 
times as broad as long, the sides and the anterior margins nearly 
straight, the posterior angles slightly oblique ; surface finely pubes- 
cent, closely and finely punctured, obsoletely depressed near the 
base and at the sides, green, opaque ; the lateral margins narrowly 
flavous, the anterior angles with an elongate narrow piceous spot. 
The apex of the scutellnm broadly truncate, the surface finely punc- 
tured, pale green. Elytra more evenly and finely punctured than 
the thorax, covered with short yellowish pubescence, the lateral 
margins narrowly flavous, the disk pale green. Underside and legs 
testaceous ; the knees, tibise, and the tarsi fuscous or piceous. Tibise 
unarmed ; claws bifid ; anterior coxal cavities open. 

A single specimen. 

Galerucella ceylonensis, sp. nov. (Plate XI. fig. 12.) 
Testaceous ; apical joints of the antennse fuscous ; head and 



10:J MR. M. JACOBY ON THE [Feb. 1, 

thorax rugose-punctate, the former with one, the latter with three 
black spots ; elytra pale fulvous, finely pubescent, a narrow sutural 
and a broad lateral stripe bright green. 

Length 3-3| lines. 

Head closely rugose-punctate, testaceous, with a large triangular 
black spot at the vertex. Antennae slender, filiform, two thirds the 
length of the body, black or fuscous at the terminal joints, the basal 
ones, varying in numbers, fulvous ; third joint longer than the 
fourth. Thorax more than twice as broad as long, the sides strongly 
rounded at the middle, tlie anterior angles produced into an acute 
point ; surface transversely and strongly depressed, more strongly 
rugose-punctate than the head, testaceous, with a large black spot 
at each side, and another one, mesially constricted, at the middle. 
Scutellum testaceous, broad, its apex truncate. Elytra slightly 
widened behind, closely covered with fine silky whitish pubescence ; 
a very narrow sutural stripe, not quite extending to the apices, and 
another very broad one, commencing at the scutellum and extending 
to the apical margin of each elytron, bright green, the rest of the 
surface as well as the extreme lateral margin testaceous. Legs of 
the same colour, the upperside of the femora and that of the tibiae 
more or less distinctly marked with a piceous stripe, sometimes the 
legs are entirely of that colour. Claws bifid. 

Kitukgalle. 

Galerucella lateralis, sp. nov. 

Ovate, slightly widened posteriorly, testaceous ; antennae, a spot 
at the vertex and three at the thorax, fuscous ; elytra closely punc- 
tured and pubescent, obscure greenish, the lateral margins narrowly 
testaceous. 

Length 2|-3 lines. 

Head finely rugose, pale brownish or testaceous, a triangular spot 
at the vertex and the central narrow groove fuscous ; the anterior 
margin of the clypeus thickened. Antennae nearly as long as the 
body, the third joint very long and slender, all the others shorter 
but equally slender, the second joint very short. Thorax more than 
three times as broad as long, the sides narrowed towards the apex, 
slightly sinuate, the anterior angles produced into a very small 
tooth, the posterior ones oblique and sinuate at each side ; the disk 
with a transverse depression at the sides and a longitudinal impressed 
central line, sculptured like the head and sparingly clothed with 
very short pubescence ; a spot at each side, and another near the 
anterior margin at the middle, fuscous. Scutellum fuscous. Elytra 
rather convex, much more distinctly and closely covered with white 
hairs, extremely finely and closely rugose-punctate, obscure dark 
greenish, opaque, the lateral margins obscure pale testaceous. The 
sides of the breast, the femora above, and the outside of the tibiae 
piceous. The first joint of the posterior tarsi scarcely longer than 
the following one ; claws bifid; anterior coxal cavities open. 

Kitukgalle. 

hi one specimen the thorax is almost entirely fuscous, owing 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 107 

probably to discoloration. At once distinguished from G. ceylonensis 
by the much more transverse and finely punctured thorax as well as 
by the uniform obscure, not metallic green, colour of the elytra. 

Galerucella crotchi, sp. nov. 

Obscure testaceous, finely pubescent ; the head with one, the 
thorax with three fuscous spots ; elytra metallic green, finely rugose 
and pubescent, the sutural and lateral margin narrowly purpHsh. 

Length 2| lines. 

Head minutely punctured, with an indistinct central longitudinal 
groove, the vertex with an obscure large fuscous spot. Antennae half 
the length of the body, testaceous, the terminal joints stained with 
fuscous, the third joint shorter than the fourth. Thorax twice as 
broad as long, the sides rounded, narrowed at the base, surface 
finely pubescent, with an obsolete transverse lateral and a deeper 
longitudinal central depression, the sides and the central groove 
fuscous. Scutellum fuscous. Elytra covered with rather long grey 
pubescence, bright metallic green, narrowly margined with purplish, 
the surface finely rugose-punctate. The tibiae unarmed, the first 
joint of the posterior tarsi nearly as long as the two following joints 
together. Claws bifid ; anterior coxal cavities open. 

Galeruca cenescens, Fairm., from Central China, seems to be closely 
allied to the present species, but diff'ers in the equal length of the 
third and fourth joints of the antennae and in the traces of longi- 
tudinal costse of the elytra, also in the different shape of the thorax. 

Galerucella marginata, sp. nov. 

Obscure testaceous ; antennae, tibiae, and tarsi black ; head and 
thorax strongly punctured, shining, the thorax with five black spots ; 
elytra very finely punctured and pubescent, obscure testaceous, a 
narrow longitudinal stripe near the lateral margin blackish. 

Length 3| lines. 

Head closely rugose-punctate, the vertex obscure fuscous. Antennae 
more than half the length of the body, black, slender, the third 
joint the longest. Thorax more than twice as broad as long ; the 
sides subangulate at the middle, the anterior angles slightly dentate; 
the surface closely impressed with larger and smaller punctures, the 
sides with a deep round fovea, the middle with an obscure longitudinal 
depression, the latter of black colour, a similar-coloured spot is placed 
close to the lateral margins, making in all five spots placed transversely 
across the disk. Elytra rather convex, very finely and closely punc- 
tured and covered with short whitish pubescence ; a narrow lateral 
stripe commences at the shoulder and is abbreviated a little distance 
from the apex of each elytron ; the first joint of the posterior tarsi 
as long as the two following joints ; claws bifid. 

A single specimen. 

Xenarthra mirabilis, sp. nov. (Plate XL fig. 9.) 

Elongate, parallel ; subdepressed, testaceous ; antennae piceous, 
pectinated, 12-jointed ; head with three, thorax with five piceous 



108 MR. M. JACOBY ON THE [Feb. I, 

spots ; elytra finely punotured, sparingly pubescent, the base and a 
broad transverse band below the middle greenish piceous. 

Length 3 lines. 

Head impunctate, a triangular spot at the middle of the vertex, 
and another smaller one at each side, piceous; eyes prominent ; the 
frontal tubercles distinct, but rather small. AntennsD as long as the 
body, piceous, the apices of all tlie joints testaceous, the first joint 
curved and slender, the second one extremely small, entirely testa- 
ceous, the eight following joints with long and slender appendages, 
the tenth much longer and broader than the preceding ones, the 
terminal joints long and slender. Thorax twice^as broad as long, 
the sides slightly rounded at, but somewhat constricted below, 
the middle, the posterior margin evenly rounded, the anterior one 
nearly straight ; the surface obsoletely transversely depressed, en- 
tirely impunctate, with a narrow longitudinal band at the sides 
and three spots, placed triangularly at the middle, piceous. Scu- 
tellum triangular, testaceous. Elytra with two deep foveas below 
the base, the punctuation rather fine and placed in close, very 
irregular rows, the interstices slightly convex and furnished with 
rows of stiff testaceous hairs ; a narrow transverse band at the base, 
the interior of the subbasilar depressions, and a broad transverse band 
below the middle, consisting of longitudinal bands joined together, 
greenish aeneous or piceous ; the elytral epipleura and the breast of 
the same colour. Tibiae sHghtly stained with piceous at their apices ; 
the latter unarmed ; the first joint of the posterior tarsi as long as 
the three following joints togetlier ; claws appendiculate ; anterior 
coxal cavities closed. 

Bogawantalawa. 

The genus Xenarthra was established by Mr. Baly on an insect 
likewise from Ceylon, and described in the ' Journal of Entomology ' 
for 18G0. The curiously shaped and deeply pectinated antennae, 
consisting of 12 or even 13 joints, will without difficulty allow the 
genus to be recognized at first sight. Closed anterior coxal cavities 
and unarmed tibiae seem to show the place of Xenarthra to be 
amongst the Plntyxanthhice of Chapuis. There is unfortunately 
only a single specimen of this handsome species before me, and being 
fixed upon a card I am not able to say with certainty to which sex 
it belongs. Mr. Baly evidently also only knew the male sex of his 
species, and it is possible that the female insect differs in the shape of 
the antennae. In the present insect a close examination of these 
parts proves them to consist of 1 3 joints, the terminal one or appendage 
being here much longer than in any other Phytophagous insect with 
which I am acquainted. 

Chapuis has described a species of Xenarthra from Abyssinia of 
which I possess a specimen ; this species, however, belongs to an 
entirely different genus. 

Xenarthra lewist, sp. nov. (Plate XI. fig. 10.) 

Entirely testaceous, the two last joints of the antennae black. 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 109 

Thorax with two deep depressions, impunctate ; elytra very finely 
and closely punctured, sparingly pubescent. 

Length 2| lines. 

Head impunctate, with a deep triangular fovea between the eyes. 
Antennae as long as the body, the third to the ninth joints furnished 
with slender and long appendages, the tenth triangularly widened and 
compressed, emarginate at its outer side, the eleventh and twelfth 
joints simple. Thorax twice as broad as long, the sides constricted 
near the base, the surface with a transverse groove at each side nearly 
extending to the middle, entirely impunctate. Scutellum triangular. 
Elytra parallel with a small depression immediately below the scu- 
tellum, extremely finely punctured, the interstices furnished here and 
there with single stiff hairs ; the first joint of the posterior tarsi as 
long as the three following joints together. 

Dikoya, 

Xenarthra unicolor, sp. nov. (Plate XL fig. 11.) 

Elongate, subdepressed ; testaceous ; the extreme apices of the 
tibiae fuscous ; thorax square-shaped, impunctate ; elytra scarcely 
visibly punctured, sparingly covered with long hairs. 

Length 4 lines. 

Head impunctate ; palpi robust. Antennae slightly shorter than 
the body, fuscous, the long appendages and the penultimate joint 
lighter.the latter thickened and elongate, as long as the following or 
terminal joint. Thorax scarcely broader than long, the sides nearly 
straight, slightly rounded before the middle ; the surface with two 
very shallow depressions, occupying the middle of the disk, clothed 
with a few long hairs, entirely impunctate. Elytra with some very 
fine punctures arranged somewhat longitudinally, clothed with long 
single hairs ; the intermediate tibiae slightly curved ; the first joint 
of the posterior tarsi as long as the three following joints together. 

Colombo. 

Of this species, which may be known by the larger size and the 
different structure of the terminal joints of the antennae, only a single 
specimen is before me. 

LUPERODES QUADRIPUSTULATUS, Motsch. 

Galle. 

This species varies in the colour of the elytral pattern from fulvous 
to black. Motschulsky's description agrees very well with the 
specimens before me, but the legs in all of them are fulvous, the 
extremities of the tibiae and the tarsi being fuscous only. Besides 
the six yellow spots of the elytra in plainly marked specimens, the 
apices of the same parts are also frequently yellow, in others the 
elytral margin is black or piceous, and the posterior yellow spot 
extends upwards at the sides, and unites with the one placed at the 
shoulder, thus reducing the yellow marks to a spot near the scutellum 
and a band at the sides, which increases greatly in width near the 
apex of each elytron. The antennae have the third joint about one 
half longer than the second, which is very short. If I have 



110 MR. M. JACOBY ON THE [Feb. 1, 

rightly referred the present species to Motschulsky's insect, the closed 
anterior coxalcavkies of L. quadripustidatus, as well as of the following 
species, would suggest rather the genus Monolepta or Nadrana for 
their reception, since the type, Luperodes alhoplagiatus, Motsch., has 
open coxal cavities. It must, however, be left for the future to 
decide with certainty the proper place of these eastern forms. 

Luperodes pectoralis, sp. nor. 

Ovate, convex ; testaceous ; the base of the head, antenna, and 
the breast black ; thorax transverse, impunctate ; elytra very finely 
punctured, testaceous, a narrow transverse band at the base and the 
extreme margins black. 

Length 2 lines. 

Head impunctate, the basal portion as well as the labrum and the 
palpi black, the rest testaceous. Antenuse as long as the body, 
entirely black, the third joint very slightly longer than the second, 
both joints short. Thorax more than twice as broad as long, the 
sides straight, the posterior margin rounded ; surface impunctate, 
without depression. Scutellum black. Elytra extremely finely and 
closely punctured, testaceous, sometimes stained with fulvous near 
the apices, the sutural and lateral extreme margins as well as a nar- 
row transverse band at the base black. Legs fulvous or testaceous, 
the tarsi obscure fuscous. Breast black ; abdomen testaceous. The 
first joint of the posterior tarsi half the length of the tibise. 

Dikoya. 

From L. basalis, Motsch., the present species diflFers in the 
entirely black antennse and the similarly coloured elytral margins, 
also in the fulvous legs. The thorax is more transversely shaped 
thau is usual in the species of this genus, but all other characters 
agree with Luperodes. 

Luperodes flavicornis, sp. nov. 

Testaceous ; breast dark fulvous ; head and thorax impunctate ; 
eljtra extremely finely and closely punctured, reddish fulvous, the 
base obscure piceous. 

Length 2 lines. 

Head impunctate, transversely grooved between the antennce ; the 
latter slightly shorter than the body, entirely flavous, the second and 
third joints short, equal ; the fourth as long or rather longer thau 
the two preceding joints together. Thorax transverse, without 
depressions, ratlier more than twice as broad as long, the sides 
slightly, the posterior margin more strongly rounded, the angles not 
produced. Scutellum obscure testaceous. Elytra slightly widened 
posteriorly, dark fulvous, very minutely and closely punctured, the 
base with a narrow transverse obscure piceous band". Posterior tibiae 
mucrouate, their metatarsus longer than the three following joints 
together. Anterior coxal cavities closed. 

The single specimen contained in this collection differs from 
L. basalis, Motsch., in the unicolorous antennae, in the differently 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. Ill 

coloured head, legs, and the want of the sutural elytral band. L. pec- 
toralis, Jac, differs in the black antennae and tarsi as well as other 
particulars. 

LUPERODES MULTIMACULATUS, Sp. nOV. 

Testaceous ; antennae and the breast black ; head with a black 
spot ; thorax minutely punctured, with a black spot at each side ; 
elytra ovate, finely punctured, the base and lateral margin anteriorly, 
a spot near the base, two others below the middle, one near the apex 
as well as the latter itself, black. 

Length 2 lines. 

Head impunctate, transversely grooved between the eyes, the 
latter large ; the vertex with a large round black spot ; labrum 
black ; antennae slender, the third joint one half longer than the 
second. Thorax transverse, rather more than twice as broad as 
long, the sides and the posterior margin rounded ; the surface 
extremely minutely punctured, testaceous, the sides with an elongate 
black spot placed close to the lateral margin. Scutellum black. 
Elytra slightly widened at the middle, extremely finely punctured 
and wrinkled, the basal and lateral margins at the anterior half as 
well as the epipleurae black ; each elytron with a small black spot 
at the shoulder, a larger and more elongate one near the scutellum, 
two similar spots placed close to each other near the suture below 
the middle (one slightly above the other), and a fifth, narrow and 
elongate spot near the lateral margin and at a little distance from 
the apex, the latter also black at the extremity. Breast black ; 
abdomen and the legs fulvous ; the metatarsus of the posterior tibias 
as long as half their length. Anterior coxal cavities closed. Elytral 
epipleurae very narrow below the middle. 

A single specimen. 

LuPERODES RUFICOLLIS, Sp. nOV. 

Black ; head and thorax rufous ; elytra black, very minutely 
punctured. 

Length H line. 

Head impunctate, bright rufous; labrum and palpi piceous. 
Antennae black, the basal joint piceous, its base testaceous, the second 
and third joints short, nearly equal. Thorax transverse, three times 
as broad as long, the sides straight, the posterior margin rather 
rounded, the surface extremely finely punctured. Scutellum black. 
Elytra exceedingly finely punctured, black. Legs slender; the 
tibiae mucronate ; the first joint of the posterior tarsi more than half 
the length of the tibiae. Anterior coxal cavities closed. 

LuPERODES ALBnpi,AGIATUS, Motsch. 

Dikoya. 

PSEUDOCOPHORA BICOLOR, Sp. nOV. 

Flavous ; thorax deeply transversely depressed ; elytra black, 
shining, obsoletely semipunctate-striate. 



112 MR. M. JACOBY ON THE [Feb. 1, 

cJ . Elytra with an oblong fovea below the scutellum, the anterior 
part of which is tuberculiform. 

2 . Elytra simple. 

Length 2 lines. 

Head impunctate ; palpi robust, the terminal joint thickened. 
Antennae filiform, entirely flavous, the third joint slightly longer than 
the following ones. Thorax transverse, the sides straight at the 
base, slightly rounded before the middle, the anterior and the 
posterior margins perfectly straight, as well as the transverse groove 
at the middle of the disk, the latter impunctate. Scutellum flavous. 
Elytra slightly widened posteriorly, the punctuation arranged in rows, 
which are more distinct anteriorly and at the sides than towards 
the apices ; the latter with a more or less distinct flavous margin or 
spot, the rest of the surface black, shining. Underside and legs 
flavous; elytral epipleune continued below the middle. Apical 
abdominal segment of the male trilobate, the intermediate lobe 
sHghtly prolonged. 

Balangoda. 

This species will enter the present genus, established by myself, on 
account of the prolonged elytral epipleurse and the punctate-striate 
elytra. The male has the elytra deeply impressed below the 
scutellum (as is the case in P. buqiietti) ; the anterior portion of this 
depression is raised in the shape of two tubercles. In P. buquetti 
these latter are placed within the fovea. The general aspect of 
P. bicolor is that of a species of Aulacophora. 

LUPERUS NIGROMARGINATUS, Sp. nOV. 

Black ; thorax obsoletely impressed, impunctate ; elytra extremely 
finely punctured, pale yellowish white, the margins narrowly black. 

Var. a. Femora pale testaceous. 

Var. b. Elytra black, the disk obsoletely paler. 

Length l|-2 lines. 

Head impunctate, transversely grooved between the eyes, the 
frontal tubercles scarcely divided. Antennae black, nearly as long as 
the body, the third joint more than twice as long as the second. 
Thorax about one half broader than long, the sides straight, the 
posterior margin slightly rounded and sinuate; surface impunctate 
or extremely finely punctured, with a depression at each side. 
Scutellum black. Elytra slightly more distinctly punctured than the 
thorax, nearly white, the sutural and lateral margins narrowly black. 
Tibiae mucronate ; the first joint of the posterior tarsi longer than 
the three following joints together. The anterior coxal cavities 
open. 

L. nigromaryinatus seems subject to a good deal of variation in 
regard to colour ; and it is probable that the form with pale elytra 
margined with black is the normal one, as even in the black specimens 
a faint paler disk of the elytra indicates the wliite portion of the type. 
In one specimen the rare instance of part colouring occurs, the left 
elytron being black, and the right one white with the black lateral 
margin. 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 113 

^NIDEA ? HIRTIPENNIS, Sp. nOV. 

Obscure testaceous ; the two apical joints of the aiiteiiuce fuscous ; 
thorax transverse, impunctate, biimpressed ; elytra very finely semi- 
punctate-striate, sparingly pubescent. 

Length 2| lines. 

Head vpith a deep fovea between the antennae, impunctate ; 
terminal joint of the i)alpi tliickened. AntenniB but slightly shorter 
than the body, fulvous, the two or three terminal joints darker, 
second joint very short, the third nearly three times as long, and 
longer than the following joints. Thorax at least twice as broad as 
long, the sides very slightly constricted at the base, a little rounded 
before the middle, the angles not produced ; the surface impunctate, 
with a rather deep oblique impression at each side. Scutellum 
triangular. Elytra without basal depression, very finely and some- 
what regularly punctured, the interstices here and there obsoletely 
raised and sparingly clothed with rather long and stiff hairs ; elytral 
epipleurae broad, continued below the middle. Tibiae unarmed ; 
the first joint of tlie posterior tarsi as long as the three following 
joints together. Claws appeadiculate. Anterior coxal cavities 
closed. 

Dikoya. 

I have placed this species in ^nidea, with which it agrees in all 
essential points. It is, however, possible that I may have only female 
specimens before me, and that the male insect, like several others of 
the genus, may differ in the structure of the head. The pubescence 
of the elytra distinguishes A. hirtipennis from any of its allies. 

CnEORANE PALLIDA, Sp. HOV. 

Oblong, pale testaceous ; antennae obscure fuscous ; thorax 
square-shaped, impunctate ; elytra scarcely visibly punctured. 

Length 2 lints. 

Head entu'ely impunctate, the frontal tubei'cles distinctly raised, 
divided and bounded behind by a deep groove ; palpi robust. 
Antennae two thirds the length of the body, the second and third 
joints short and of nearly equal length, the fourth joint very slightly 
longer than the following ones, the three or four basal joints pale 
testaceous, the rest fuscous. Thorax shghtly broader than long, all 
the margins straight ; the surface very little convex, without depres- 
sions and punctures. Elytra rather convex, parallel and sub- 
cylindrical, the punctuation extremely minute and arranged for the 
greater part in closely placed rows. The tibiae unarmed, the first 
joint of the posterior tarsi as long as the two following joints 
together. Claws appendiculate. Anterior coxal cavities open. 

Bogavvantalawa. 

In this species the third joint of the antennfe is sliorter than in 
C. elegans, Baly, and Cfulvicollis, Baly, being of the same length 
as the second joint ; but in all other respects C. pallida agrees with 
its allies, and may be recognized by the uniform and very pale 
testaceous colour. 

Proc. Zool. Soc— 1887, No. VIII. 8 



1!4 MR. M. JACOBY ON THE [Feb. 1, 

Mktrioidea rufipennis, sp. nov. (Plate XI. fig. 8.) 

Oblong ; black ; head, anteuuse, and legs fulvous ; thorax greenish 
black, obsoletely depressed ; elytra rufous, very finely punctured. 

d . Face deeply excavated ; the third joint of the antennas curved 
and produced at the apex. 

Length .S lines. 

Head rufous at the vertex, impunctate, deeply transversely grooved 
between the eyes ; lower part of the face deeply excavated, the 
excavation bounded at the sides and above by several lobes which 
protrude beyond the impressed portion ; the clypeus thickened ; 
palpi moderately incrassate at the penultimate joint. Antennae 
nearly as long as the body, fulvous, the second joint very short, the 
third curved and widened into a tooth at the apes, nearly as long as 
the following joints, the apical joints more slender than the rest. 
Thorax about one half broader than long, narrowed towards the base, 
greenish black, shining, the surface with an obsolete transverse 
depression at the sides near the base, impunctate. Scutellum rufous. 
Elytra very finely and rather closely punctured, rufous, their 
epipleurae continued below the middle ; tibise unarmed, the first 
joint of the posterior tarsi as long as the two following joints 
together. Claws appendiculate The anterior coxal cavities closed. 

Kandy. 

1 have placed this species, of which I have evidently only the male 
insect before me, in the present genus on account of the closed coxal 
cavities, unarmed tibise, and the appendiculate claws ; the propor- 
tionate length of the joints of the antennse is, however, different than 
in Metrioidea, and it is possible that the present species is repre- 
sentative of a new genus. 

OCHRALEA CEYLONICA, Harold (?). 

Dikoya. 

The descriptions of two species of Ochralea from Ceylon have 
been piiblisbed by von Harold. With one of these the insect which 
I refer to the present species agrees in the main points. It is, 
however, smaller by one millimetre ; the antennse, with the exception 
of the two basal joints are fuscous, not testaceous, the third joint 
i)eing slightly longer than the second. The elytra have the sides 
more or less stained with obscure fulvous, and the punctuation is 
exceedingly close, and consists of larger and smaller punctures. 
Without examining the type of O. ceylonica contained in the Berlin 
Museum, it is impossible to say whether the specimens before me 
represent that or an allied species. In regard to the genus Ochralea, 
Mr. Baly has drawn my attention to the state of the anterior coxal 
cavities, which according to Chapuis are supposed to be closed. A 
careful examination of several specimens proves this, however, to be 
erroneous, as the cavities are distinctly open. This character and the 
prolonged elytral epipleurse will not allow Ochralea, according to 
Mr. Baly's opinion, to be separated from Luperodes, a genus which 
seems also to possess open cavities, although I have considered the 
latter in Luperodes as being closed. The whole question of open or 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEVLON. 115 

closed coxal cavities requires yet careful study, as it is not improbable 
that intermediate degrees exist even in the same species, which makes 
the conclusion in regard to the state of the cavities uncertain. 

Hyph^nia flavofemoratus, Motscli. (?). 

Closed anterior coxal cavities, unarmed tibise, a square-shaped 
thorax, and other characters peculiar to Baly's genus are present in 
an insect contained in this collection, which also agrees very nearly 
with Motschulsky's species, to which I will refer for the present this 
insect. The entire upper surface is metallic greenish or seneous. The 
antennae are as long as the body, obscure piceous with the basal 
joints fulvous. The head is finely granulate, the lower part being 
testaceous ; the thorax is square-shaped, with two impressions, finely 
granulate and punctured (Motschulsky gives the thorax as smooth). 
The elytra are finely rugose and punctured. The colour of the legs 
is flavous ; sometimes the tibise and the tarsi are obscurely stained 
with fuscous. The femora are all of the same thickness, and the first 
joint of the posterior tarsi is as long as the two following joints 
together. Claws appendiculate. The underside is nearly black ; and 
the entire shape of the insect is narrowly parallel with the elytra 
flattened along the sutural margin. The size of the insect is 1| line. 

Dikoya. 

DoRYScus (gen. nov. Galerucinse). 

Body elongate, parallel ; anterior coxal cavities closed. Antennas 
filiform, the second joint small, the third double the length. Thorax 
subquadrate, strongly constricted at the base, the disk biimpressed. 
Elytra geminate punctate-striate, costate, and pubescent. Tibise 
mucronate ; the first joint of the posterior tarsi as long as the two 
following joints together ; claws appendiculate, those of the posterior 
tarsi very long and curved, united, but bifid at the extreme apices. 
Elytral epipleurse narrow, but continued below the middle. 

Whether the peculiar structure of the posterior claws in this genus 
is a sexual character only or to be found in either sex I am unable 
to say, having only two specimens before me, which agree with each 
other in every respect. The posterior claw seems to consist of a 
single piece only, being joined together except at the extreme apices, 
as is the case, but to a much smaller degree, in the genus Lema. In 
Doryscus they are very long and curved, while the claws of the four 
anterior legs are of normal size and appendiculate. The peculiar 
form of the thorax, in connection with the punctate-striate elytra and 
their pubesence, are characters which further distinguish the present 
insect. 

Doryscus testaceus, sp. nov. 

Testaceous ; the sutural and extreme lateral margin narrowly black 
anteriorly; head and thorax impunctate; elytra strongly geminate 
punctate-striate, the interstices longitudinally costate. 

Var. Entirely testaceous. 

Length 2 lines. 

8* 



116 MR. M. JACOBY ON THE [Feb. 1, 

Head impunctate, the frontal tubercles rather flattened ; palpi 
slender ; antenuEe two thirds the length of the body, testaceous, the 
apical joints obscure fuscous. Thorax about one half broader than 
long, the sides strongly narrowed or constricted at the base, 
obsoletely impressed at the sides, impunctate, the lateral margins 
furnished with some long hairs. Scutellum broader than long, 
testaceous or margined with piceous. Elytra clothed with some 
rather long and stiff hairs, strongly longitudinally costate at the 
sides, less strongly at the disk, the interstices impressed with two 
rows of distinct punctures, partly confluent and becoming single 
towards the apices. Underside testaceous, the breast sometimes 
darker. Legs moderately long, the claw-joint of the posterior tarsi 
very elongate, as long as the metatarsus. 

Priapina (gen. nov. Galerucinae). 

Narrowly oblong. Antennae filiform, the third joint very small. 
Thorax transversely subquadrate, obsoletely grooved at the disk. 
Elytra closely semipunctate-striate. Legs slender, tibiae with a 
spine ; the metatarsus of the posterior tibise longer than the three 
following joints together ; claws appendiculate. 

I am obliged to establish this genus on account of the structure 
of the antenuse, in which the third joint of the male is so small as 
to be scarcely perceptible ; the long first joint of the posterior tarsi 
and the mucrouate tibise are further characters which will assist in 
the recognition of Priapina. The specimen being glued to a card, 
I am unfortunately not able to speak about the state of the anterior 
coxal cavities. There will, however, be no difficulty in recognizing 
the genus if the antennae are examined. Li Luperodes, & closely 
allied genus, the third joint in both sexes is very distinct, double the 
length of the second and scarcely shorter than the fourth. 

Priapina longicornis, sp. nov. 

Fulvous or testaceous ; head impunctate ; terminal joints of the 
antennae fuscous ; thorax fulvous, rugosely punctured; elytra testa- 
ceous, the margins narrowly piceous, surface closely punctured. 

<S . Antennae longer than the body, the third joint minute. 

2 . Antennae slightly shorter, the third joint a little longer ; 
elytra coarsely punctured. 

Length 1 line. 

cS . Head impunctate, fulvous, the frontal tubercles narrowly trans- 
verse ; eyes rather large. The four lower joints of the antennae 
testaceous, the others fuscous, the fourth joint longer than the three 
preceding ones together and longer than the following joints. Thorax 
twice as broad as long, the sides slightly narrowed towards the base, 
the angles not produced, the anterior and posterior margins straight ; 
surface obsoletely transversely depressed, coarsely punctured. 
Scutellum piceous. Elytra testaceous, narrowly margined with 
pioeous, very closely punctured, the punctuation arranged in semi- 
irregular rows. Legs rather long and slender. 

In the female the third joint of the antennae is nearly equal in 



1887.] PHYTOPHAGOUS COLEOPTERA OF CEYLON. 117 

length to the second, but the fourth joint, as in the male, is the longest, 
and the elytra are coarsely punctured ; the general size is also larger 
and more robust. 
Dikoya. 

Neochrolea (gen. nov. Galerucinse). 

Body oblong. Head longer titan broad, the front excavated above 
and below the antennae ; the latter as long as the body, filiform, the 
second joint very small, the third the longest, thickened and 
emarginate below. Palpi thickened at the penultimate joint. Thorax 
transversely subquadrate, the disk obsoletely depressed. Elytra 
scarcely visibly punctured, their epipleurse continued below the 
middle. Tibiae mucronate. The first joint of the posterior tarsi a 
long as the two following joints together. Claws appendiculate 
Anterior coxal cavities closed. 

Neochrolea seems allied to the genus Macrima, Baly, with which 
it agrees in the closed coxal cavities and the mucronate tibia) ; it 
differs in the long third joint of the antennae and in the less transverse 
thorax, also in the shorter first joint of the posterior tarsi. The 
only specimen before me is evidently a male ; and it is probable that 
the female wants the deep excavations of the head, as is often the 
case in similar structures in the sexes in other genera. yEnidea, 
Baly, differs in the unarmed tibiae. 

Neochrolea cavifrons, sp. nov. (Plate XI. fig. 4.) 

Entirely testaceous ; third joint of the anlenna3 emarginate below ; 
Lead with a deep excavation; thorax nearly impunctate, the disk 
obsoletely depressed ; el} tra extremely finely and closely punctured. 

Length 4 lines 

Head longer than broad, impunctate, the space below the antennae 
deeply excavated, the lower margin of this excavation forming a 
triangular and pointed flattened projrction ; the lower part ot the 
face again deeply excavated ; the anterior margin of the clypeus 
produced into two long points ; the extreme apices of the jaws 
black ; palpi thickened at the penultimate joint. Antennae slightly 
longer than the body, fulvous, the second joint extremely short ; the 
third elongate, thickened, and hollowed out at the lower margin ; the 
following joints of half the length, equal. Thorax one half broader 
than long, narrowed towards the base, the sides very little rounded 
before the middle ; the surface with a small dejjression at the middle 
of the disk, scarcely visibly punctured. Elytra rather convex, 
without basal depression, not more distinctly punctured than the 
thorax, testaceous like the rest of the insect. 

Balangoda. 

Haplotia (gen. nov. Galerucinse). 

Narrowly elongate. Antennae 'slender, filiform, all the joints 
with the exception of the second of nearly equal length ; palpi with 
the penultimate joint thickened. Thorax square-shaped, obsoletely 
impressed, rugose. Elytra closely rugose (cj), or simply punctured 



118 ON PHYTOPHAGOUS COLEOPTERA OF CEYLON. [Feb. 1, 

( 2 ) ; their epipleurse continued below the middle. Legs slender and 
elongate ; the tibiae unarmed ; the first joint of the posterior tarsi 
longer than the two following joints together ; claws appendiculate. 
Prosternum invisil)le. The anterior coxal cavities closed. 

The insect for the reception of which I am obliged to establish 
this genus seems allied to Metrioidea, Fairm., on account of the 
closed cavities and unarmed tibiae ; but differs in the proportionate 
length of the joints of the antennae, the third joint in Metrioidea 
being described as a little longer than the second and shorter than 
the fourth ; the first joint of the posterior tarsi is also longer than in 
Metrioidea. In the insect before me the female, on account of its 
different coloration, seems at first sight to constitute a different 
species. The general appearance of the present species is that of a 
Luperus, 

Haplotia VARiPENNis, sp. Dov. (Plate XI. figs. 5, 6.) 

S . ^neous ; the base of the femora and tibiae and the abdomen 
testaceous ; head finely punctured in front ; thorax and elytra 
rugosely punctate. 

$ . Testaceous, head and thorax aeneous ; elytra irregularly 
punctured, testaceous with metallic gloss ; a triangular s|)ace at the 
base, surrounding the scutellum, and the lateral margin aeneous. 

Length l|-2 lines. 

Head broad, finely rugose at the anterior portion ; labrum obscure 
fidvous. Antennae a little shorter than the length of the body, 
black in the male, obscure fulvous in the female. Thorax square- 
shaped, verv sli^'htlv narrowed at the base, tlie anterior angles 
acute, the posterior ones obsolete ; the surface flattened, closely and 
irregularly rugose, of greenish bronzed colour like the head. Elytra 
of the same colour, sculptured like the thorax, sparingly covered 
with some stiff hairs. Legs more or less piceous or aeneous, the 
base of the femora often testaceous, the posterior femora in the male 
extending to the apices of the elytra, but much shorter in the 
female. 

Nuwara Eliya. 

The antennae and the legs in the female are generally of a dark 
fulvous colour ; the entire underside is testaceous, or sometimes 
slightly stained with aeneous ; the sculpture of the head and thorax 
agrees with that of the male. The elytra are, however, not rugose 
or very slightly so, but generally closely punctured and of a pale 
testaceous "colour, slightly tinged with metallic greenish ; the base has 
a well-defined trian-iular" spot, extending from the shoulder to the 
tuture, of metallic bronze colour, the lateral margin and apices of 
the elytra being similarly coloured. Some specimens show a small 
testaceous lateral margin of the thorax, and two more or less distinct 
depressions at the disk of the latter. 



Antipha NiETNERi, Baly. (Plate XI. fig. 7.) 

Balangoda. 

Typical and unicolorous unspotted forms. 



,^^e^^sHl7(}^^ 



/N « 



=fe 



•^L H18l°^'' 



p. Z,S, 1887.. PL. XII. 




J.Srnit Lin. 



lianhaiL irnji. 



BRACHYUP.US CALVUS 



1887.] MR. F. E. BEDDARD ON BRACHYURUS CALVUS. 1 19 

EXPLANATION OP THE PLATES. 
Plate X. 

Fig. 1. Demoiina semifasciata, p. 70. 

2. Pagria costatipennis, p. 73. 

3. Ehyparida qidnqicemaculata, p. 75. 

4. Nodostoma tiihcrosum, p. 78. 

5. Dennorrhytis igneofasciata, p. 83. 

6. ornatissima, p. 82. 

7. ceylonensis, p. 8L 

8. Chabria nigraplagiafa, p. 93. 

9. apioicornis, p. 93. 

10. Pexodorus ceylonensis, p. 95. 

11. Ivalia metallica, p. 100. 

12. viridipennis, p. 100. 

Plate XL 

Fig. 1. Aidacophora stevensi, p. 103. 
2, 3. nigripeta, p. 103. 

4. Neochrolea cavifrons, p. 117. 

5, 6. Haplotia varipennis, p. 118. 

7. Antipha nietneri, p. 118. 

8. Metrioidea riifipeyinis, p. 114. 

9. Xenarthra mirabilis, p. 107. 

10. ?et«isj, p. 108. 

11. unicolor, p. 109. 

12. Galerucella ceylonensis, p. 105. 



3. Notes on Brachyurus calvus. By Frank E. Beddard, 
M.A., F.R.S.E., Prosector to the Society. 

[Eeceived January 13, 1887-] 
(Plate XII.) 

The accompanying drawing (Plate XII.) represents the external 
characters of the male Brachyurus calvus which died in the Society's 
Gardens on July 2 1 last year. I have taken the opportunity of com- 
paring the structure of this species with the closely-allied B. rubi- 
cundus, which has been carefully described by Forbes in his memoir 
on the Ouakari Monkeys'. 

The genus Brachyurus contains three species ^ of which two, 
viz. B. calvus and B. rubicundus, agree very closely in external 
characters, and, as I shall presently show, in internal structure ; 
while the third, B. melanocephalus, differs more in external characters 
from the other two than they do from each other. 

Mr. Forbes has given a detailed account of the external characters 
of B. rubicundus, and the main external characters of all the species 
are referred to by Schlegel ^. The general coloration of the back is a 
whitish grey, produced by a mixture of white and black hairs, the 
white predominating ; passing from the dorsal to the ventral surface 

' P. Z. S. 1880, p. 627. 

^ Schlegel' " Pithecia alba " was believed by Mr. Forbes to be identical with 
B. calvus. 

^ Museum des Pays-Bas, 1876, p. 227 et seq. 



120 MR. F. E. BEDDARD ON BRACHYURUS CALVUS. [Feb. 1, 

the colour gradually assumes a fulvous-brown tint, the brown being 
darker in the pectoral region, where the brown hairs are very 
numerous, only a few white hairs being interspersed among them. 
The brownish tinge is also conspicuous on the amis, Itgs, and tail, 
particularly on the tail, on the jjosterior aspect of the thighs, and at 
the wrist and ankle. The top of the head is a greyish colour, 
gradually passing into brown anteriorly and at the sides, as in 
B. ruhkv7idus ; the hairs on the throat also resemble that species in 
their dark brown colour, and in being mixed with numerous black 
hairs ; the general tint of the hair on the throat is a rich chestnut- 
brown, and is exactly similar to that of B. rubicundus. 

With regard to the osteology, I find that the number of vertebrae 
in my specimen is C. 7, D. 13, L. G, S. 4, Cd. 15, of which the 
last three are very minute and apparently ankylosed together. 
Forbes states in his paper that in B. melanocephalus there are 19 or 
20 caudal vertebrse, on the authority of a specimen belonging to 
that species in the National Collection. The specimen in question 
(806 h) has certainly the 20 caudal vertebras that Forbes has men- 
tioned ; but it does not present any recognizable differences from 
Brachyurus calvus, and indeed is entered in the Catalogue as belonging 
to that species. 

It is not necessary to give much account of the visceral anatomy 
of this species, inasmuch as I have been unable to find any marked 
points of difference from B. rubicundus ; the alimentary viscera pre- 
sented a very close correspondence in the two species, as will be 
evident from the following notes. 

The tongue resembles in every particular that of B. rubicundus, 
and, curiously enough, even the arrangement of the circumvallate 
papillae corresponds in the two species. The corresj)ondence is curious, 
because Mr. Forbes's description of the circumvallate papillae reads 
almost as if he were referring to an abnornal condition. The circum- 
vallate papillae in the two species are disposed in the usual V-shape, 
but there is an additional papilla on the right side between the apical 
and basal papillae, thus destroying the symmetry of the arrangement. 
In a specimen of Macacus rhesus, to which I am able to refer at the 
moment of writing, there are also four circumvallate papillae ; two are 
situated side by side, and symmetrically at the apex of the V, while 
the two others occnjiy the usual position. 

Cacum. — The caecum measured 10 inches along the greater cur- 
vature ; it is separated from the colon by a very marked constriction; 
it is not sacculated, and when fully distended with air was curved 
on itself into a little less than a circle ; it is furnished with a well- 
developed median frenum which carries blood-vessels. 

In examples of two species of Cullilhrix and in a Pithecia I have 
noted an identical structure in the caecum. 

The origin of this peritoneal fold is not exactly in the middle line 
at the lower extremity of the ileum, and the blood-vessel passes on to 
it over one side of the base of the ileum ; the blood-vessel in fact 
exactly corresponds to that which is borne by one of the lateral folds 
in Hapale. 



1887.] MR. THOMAS ON MAMMALS FROM THE CAMEROONS. 121 

In Ilapale jacchus the cseoum distended with air, dried and var- 
nished, showed three folds of peritoneum running along its ujiper 
surface, as described by Prof. Flower' in Ateles ; the fremtm or 
median band is extremely short and bears no blood-vessel. The 
lateral folds arise precisely as is indicated by Prof. Flower, but one of 
them is much longer than the other and reaches nearly to the end of 
the caecum, while the other does not reach so far as does the median 
frenum. 

In Midas rufimanus a spirit-specimen of the caecum showed the 
same three folds, which were, however, jiartially united together into 
an apparently single fold ; this was easily separable into three layers 
— a median fold without blood-vessels, and two lateral folds, each 
bearing a blood-vessel. 



4. List of Mammals from the Cameroons Mountain^ collected 
by Mr. H. H. Johnston. ^. By Oldfield Thomas. 

[Eeceived January 4 ,1887.] 

In order to complete the list of the zoological specimens collected 
by Mr. II. H. Johnston, I have been asked to contribute the names 
of the two Mammals he obtained. They are as follows : — 

1. Anomalurus BEECROFTi, Fraser. 

a. Skin and skeleton, c? . Cameroons Mountain, 8000 feet. 

2. Mus UNiviTTATUs, Pcters. 

a. Skin, 5 • Cameroons Mountain, 8000 feet. 

1 Med. Times and Gazette, 1872. 

^ [Mr. Johnston's narrative of his ascent of the Cameroons Moxmtain last year, 
during v\hich the collections described in this and the folloTving communications 
■were made, w ill shortly appear in the ' Graphic ' with illustrations. Setting out 
from Victoria, opposite hi.=! residence on Mondole Island, Mr. Johnston pro- 
ceeded by Eonjougo and Mapanja (3000 feet alt.) to Mann's Spring, where lie 
encamped at an altitude of 7-Jf feet. Here the temperature ranged from 50° 
to 60° Fahr., and for the first week of his stay he lived in a perpetual rainfall. 
The forest-region ceases at about 7000 feet, and gives jilace to grassy downs, 
dotted with patches of woodland and varied by huge isolated boulders of i-ock 
and ancient lava-flows. Here a corresponding change in the flora and fauna 
takes place. Mr. Johnston tells us : — 

"Mann's Spring is a favourite resort of birds, who alway affect the vicinity of 
water, and here especially they make the air musical with their twittering songs 
and mellow love-calls. As man is a rare visitant here, the birds are very bold 
and fearless, and appeared to welcome our coming for the chance scraps of food 
thrown in their way. Alas ! they soon had to rue their over-confidence. They 
had put themselves in the power of one whose natural tender-heartedness and 
love of living things are overborne by his interest in science. Of all the pretty 
bird-forms which came to drink and sport and bathe by the brooklet, or which 
hovered about the balsam-blossoms, some of every kind must die to illustrate 
the ornithology of the Cameroons. And so my native collector and I were soon 



122 CAPT. G. E. SHELLEY ON [Feb. 1, 



5. On a Collection of Birds made by Mr. H. H. Johnston 
on the Cameroons Mountain. By Captain G. E. 
Shelley^ F.Z.S. 

[Eeceived January 3, 1887.] 
(Plates XIII. & XIV.) 

Mr. H. H. Johnston, F.Z.S., well known for his researches on 
the Congo, and successful expedition to the heights of Kilimanjaro 
in East Africa, has now sent us some birds from a nearly equally 
elevated district of Western Africa ; and I am pleased to find in this 
collection from the Cameroons Mountain an interesting proportion 
of new species. The collection, which has been sent to me for 
examination by the Cameroons Committee of the British Association, 
contains 36 skins referable to 18 species. Of these the following are 
new to science : — (1) Poliopicus Johns font, (2) Psalidoprocne fidigi- 
nosa, (3) Laniarius atroflavus, and (4) Ploceus melanog aster. 

Our previous knowledge of the avifauna of the higher part of the 
Cameroons Mountain is entirely derived from an article by Mr. G. R. 
Gray (Ann. Nat. Hist. 1862, x. p. 413) on the birds obtained by 
Capt. R. Burton during his ascent of the mountains in 1861-62'. 
In 1871 Mr. R. B. Sharpe (P. Z. S. 1871, p. 614) described a 
collection of birds made by Mr. Crossley in the Cameroons district ; 
and in 1874 and 1875 Dr. Reichenow, in the ' Journal fiir Orni- 
thologle,' published the results of his West- African Expedition of 
1872, during which he visited the Cameroons river and penetrated 
up the mountain to a height of about 4000 feet. But neither 
Mr. Crossley nor Dr. Reichenow seemed to have obtained any 
specimens from the higher elevations to which Capt. Burton and 
Mr. Johnston have ascended. 

1. POLIOPICUS JOHNSTONI, Sp. n. 

a. S , October, 6000 feet. — A broad black forehead with a buff 
patch on each side of the base of the culmen ; remainder of the 
crown and the nape red. Remainder of the upper parts, when the 

engaged in skinning black and golden Shrikes, metallic-green and crimson- 
breasted Sunbirds, ruddy Chats, olive-green Warblers, dull grey Grosbeaks, and 
tiny, indefinite, insect-eating birds of blue-grey and russet-brown. 

" In this forest, too, I shot flying Squirrels and small vole-like Eats. These 
were the only mammals we saw, except when, very rarely, we got a hurried 
glimpse of a red-coated, white-.stripcd Antelope of the genus Tragelaphu^." 

From Mann's Spring Mr. Johnston transferred his camp to Hunter's Hut 
(8300 feet), in " a narrow peninsida of forest which pushes up the mountain- 
side," and subsequently to another spot situated at an elevation of 10,.)00 feet, 
whence the final ascent was made. He made the summit by boiliug-jDoiut 
observation to be 13,760 feet, which is about 500 feet less than the usual 
estimation, — P. L. S.] 

^ See Burton's ' Abeokuta and the Cameroons Mountains.' 2 vols. 8vo. 
London, 1863. 




en 

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P.Z.S,1887. PI. XIV. 




J.&Kp.uletTidn& ttli . 



Haiihavt imp . 



Fiq ], ZOSTEROPS MELANOGEPHALA 



'3 



2.PL0CEUS MELA^QGASTFR 



1887.] BIRDS FROM THE CAMEROONS. 123 

wings are closed, uniform olive-green, with the primaries and tail 
dark brown, slightly washed on the edges of the feathers with 
yellowish olive. The shafts of the quills and tail-feathers are brown 
above and yellow beneath. Sides of the head huffish olive. Under 
surface of the body uniform sulphur-yellow, with a very slight 
greenish shade, paler on the throat and fading almost into white on 
the chin. A few feathers on the sides of the lower throat show 
faint signs of dark shaft-stripes, which shaft-stripes become more 
strongly marked on the flanks and under tail-coverts. Under wing- 
coverts yellowish buff; under surface of the quills dark brown with 
pale yellowish shafts, and with from two to four large yellowish-buff 
spots on their inner webs, giving a barred appearance. Tail beneath 
more olive than above. Bill whitish, becoming dark towards the 
base ; legs and feet dusky brown. Total length 6-6 inches, culmen 
0-8, wing 3-45, tail 2-8, tarsus 07. 

b. c?, 6000 ft. — Perfectly similar in plumage to a. Total 
length 6-3 inches, culmen 0*7, wing 3*5, tail 27, tarsus 07. 

This bird, which is closely allied to P. ellioti from the Gaboon 
and Congo district, is rather smaller. Its chief character, which 
shows no variation in Mr. H. H. Johnston's two specimens, is the 
almost entire absence of markings on the underparts, which 
parts in P. ellioti are strongly striped with brownish black in both 
sexes. 

2. Indicator variegatus, Less. 

Indicator variegatus, Reichen. J. f. O. 1875, p. 6, Cameroons ; 
Sharpe in Dawson Rowley's Orn. Misc. i. p. 189. 
$ , September, 7000 ft. 

3. Corythaix meriani, Riipp. 

Oorythaix meriani, Hartl. Orn. W.-Afr. p. 157 ; Sharpe, P. Z. S. 
1871, p. 605, Cameroons; Schalow, J. f. O. 1886, p. '67. 
S , October, 5000 ft. 

4. PSALIDOPROCNE FULIGINOSA, Sp. n. 

c? , September, 9000 ft. — Entire plumage dark brown with no 
gloss. Quills, tail, and underparts slightly darker. Under wing- 
coverts brown, scarcely paler than the back. Bill black ; feet brown. 
Total length from tip of bill to tip of tail 5 inches, culmen 0-2, wing 
4-25, tail 3"1, tarsus 0-35. 

$ , September, 9000 ft. — Perfectly similar in plumage, with the 
outer web of the first primary equally serrated. Total length 5 
inches, culmen 2, wing 4-1, tail 27, tarsus 0-35. 

In this species the tail reaches about to the tips of the wings but 
not further, and is only moderately forked ; length of centre feathers 
2-3 and 2 inches, of outer feathers 3 and 27. 

It is probably nearest to P. jietiti, from which it differs in the 
colour of the under wing-coverts, and the moderately forked tail not 
extending beyond the tips of the wings. 



124 CAPT. G. E. SHELLEY ON [Feb. 1, 

5. Trochocercits, sp. ? 

? Trochocercus nitens, Cass. Pr. Philad. Acad. 1S59, p. 50 ; Sharpe, 
Cat. B. Brit. Mus. iv. p. 300. 

? Terpsiphone nigromitrata, Reiclien. J. f. O. 18/4, p. 110 ; 1875, 
p. 24, Cameroons, 

5 , September, 7000 ft. 

The poor condition of this specimen prevents me from confidently 
determining what name really belongs to it. It is in all probability 
2\ nigromitratus, which was found in the Cameroons, and which title 
Mr. Sharpe refers to T. nitens, Cass. 

The following; is a descrijjtion of Mr. Johnston's specimen : — 
There is no gloss on the plumage. Upper half of the head black 
and but slightly crested, neck and back dusky slate-colour ; wings 
and tail black, with scarcely any signs of grey edging to the feathers. 
Sides of the head and upper throat dusky grey, nearly black ; lower 
crop, throat, and flanks slate-colour; remainder of the under surface 
of the body white, shading into rufous buff on the under tail-coverts ; 
thighs brownish slate-colour. Bill black ; legs dark brown. Total 
length 4"8 inches, culmen 0'4, wing 2*45, tail '2'7, tarsus 0'65. 

6. Laniarius atroflavus, sp. n. (Plate XIII.) 

(5 , October, 7300 ft. — Upper parts glossy black ; feathers of the 
lower back fluffy, and some of them with a large rounded white sub- 
terminal spot more or less hidden by the overlying black feathers ; 
the last feathers of the rump with broad huffish ends, forming a 
band at the base of the tail. Underparts deep yellow, paler on the 
upper half of the throat and chin, changing into rufous buff 
between the thighs and on the under tail-coverts ; outside of the 
thighs black ; axillaries yellow, under wing-coverts black. Bill and 
legs black. Total length 7 inches, culmen 0*7, wing 3'3, tail 3'2, 
tarsus 1'2. 

2 , October, 7300 ft.- — Differs only from the male in the flanks 
being dusky black. Total length 6'7 inches, culmen O'S, wing 3'2, 
tail 3'1, tarsus 1"2. 

This species is, I think, nearest to L. sublacteus (Cass.), which 
chiefly differs in having the entire underparts white. 

7. Xenocichla tephrol^ma (Gray). 

Trichopho7-us tephrolcemvs. Gray, Ann. Nat. Hist. 1862, x. p. 444. 

Xenocichla tephrolcema, Sharpe, Cat. B. Brit. Mus. iv. p. 98. 

2, September, 7000 ft. ; 2, October, 7000 ft. 

The type specimen was killed by Capt. R. Burton in the Cameroons 
Mountain at 7000 ft., so that the species would appear to be very 
local. 

8, Callene ISABELLA (Gray). 

Cossypha isabellce. Gray, Ann. & Mng. Nat. Hist. 1862, x. 
p. 443. 

Callene isabellce, Sharpe, Cat. B. Brit. Mus. vii. p. 17. 
6, September, 7000 ft.; d, October, 7000 ft. 



1887.] BIRDS FROM THE CAMEROONS. 125 

This is another apparently very local species, also procured by 
Capt. R. Burton in the Cameroons Mountain at 7000 ft. 

9. Pratincola axillaris, Shelley. 

Pratincola axillaris, Shelley, P. Z. S. 1884, p. 556 ; 1885, 
p. 220. 

cJ , September, 8000 ft. ; S , October, 8000 ft. 

This is the second species of Pratincola recorded from the 
Cameroons ; for the P. salax, Verr., of Gray's List, two specimens of 
which I have examined in the British Museum, has the axillaries 
and under wing-coverts white. 



'o 



10. ZosTEROPS melanocephala, Gray. (Plate XIV. fig. 1.) 
Zosterops (Speirops) melanocephalus, Gray, Ann. & Mag. Nat. 

Hist. 1862, X. p. 444. 

5 $ , September, 7000 and 8000 ft. 

The type, which I have examined in the British Museum, came 
from 7000 ft. in the Cameroons Mountain. 

11. Cisticola ruficapilla (Fraser). 

Brymoica i-uficapilJa, Haiti. Orn. W.-Afr. p. 57 ; Reichen. 
J. f. O. 1875, p. 45, Cameroons. 

Cisticola ruficapilla, Sharpe, Cat. B. Brit. Mus. vii. p. 248. 
a. ? , September, 7000 ft. ; b. 8000 ft. 

12. CiNNYRis CHALYBEUS (Linn.). 

Cinnyris chalyheus, Shelley, IVIonogr, Sun-birds, p. 253, pi. 78. 

cJ, September, 8000 ft. Length of wing 2*2 inches, culnien 0"8. 

cJ, September, 7000 ft. Length of wing 23 inches, culmen 0'8. 

2 , September, 7300 ft. Length of wing 2'0 inches, culmen 0'7. 
The males have the abdomen, under tail-coverts, and under surface 
of the tail darker than in any South-African specimens I have seen, 
but all the other characters are perfectly similar. This is the first 
time the species has been recorded from the West-African sub- 
region. 

13. Cinnyris fuliginosus (Shaw). 

Nectarinia fuliginosa, Hartl. Orn. W.-Afr. p. 43; Sharpe, P. Z. S. 
1871, p. 30, Cameroons; Reichen. J. f. O. 187.5, p. 30. 

Cinnyris fuliginosus, Shelley, Monogr. Sun-birds, p. 275, pi. 86. 
a. Not labelled. 

14. Anthus pyrrhonotus (Vieill.). 

Anthus gouldii, Hartl. Orn. W.-Afr. p. 73. 
Anthus pyrrhonotus, Sharpe, Cat. B. Brit. Mus. x. p. 555. 
S, October, 1000 ft. Length of wing 3-4. 
Mr. Sharpe writes {loc. cit.) : — " In Western Africa a small dark 



126 CAPT. SHELLEY ON BIRDS FROM THE CAMEROONS. [Feb. 1, 

race occurs from the Niger to Senegambia, while the representative 
form of the Gaboon and Congo region is subspecifically distinct." 
Mr. Johnston's specimen I consider belongs to the " small dark 
race." 

15. COLITJSPASSER CAPENSIS (Limi.). 

Euplectes phcenicomerus. Gray, Ann. & Mag. Nat. Hist. 1862, x. 
p. 444, Cameroons. 

CoUuspasser capensis, Shelley, Ibis, 1885, p. 359. 

S ad., October, 11,200 ft.; " d ad., 10,000 ft. ; c? ad., 8000 ft. ; 
$ , September, and d"juv., October, 8000 ft.; $ , October, 10,000 ft. ; 
cJ juv., September, 9000 ft. 

All these specimens belong to the small South-African race of 
C, capensis, which was separated by Dr. Cabanis under the name 
of Orynx approximans. 

I wish here to correct an error I made in last year's 'Ibis,' p. 350, 
where I included W. Africa in the range of the closely allied C. 
xanthomelas , It has never yet been found on that side of the 
continent. 

16. Ploceus melanogaster, sp. n. (Plate XIV. fig. 2.) 

S , September, 8000 ft. 

Head and entire throat bright yellow ; a band through the eye, 
sides of the neck, entire body, wings, and tail black. Total length 5 
inches, culmen 0'65, wing 2-6, tail 2*2, tarsus 0'8. 

The bill is comparatively long and slender. The whole bird so 
closely resembles in size and form P. nigricollis, Vieill., that I expect 
the female will be found to have a black crown. 
. This very distinct species may at once be recognized by its 
entirely black body and yellow throat. 

17. Crithagra BX3RTONI (Gray). 

Strolilophaga hurtoni. Gray, Ann. & Mag. Nat. Hist. 1862, x. 
p. 445, Cameroons. 

c? c?, October, 9000 ft. ; $, 9000 ft. 

As this is the first time the male has been sent to this country, I 
would observe that it is similar in plumage to the female. 

18. Sterna macrura, Naum. 

Sterna brachypus, Hartl. Orn. W.-Afr. p. 255. 
Sterna hirundo, Dresser, B. Eur. viii. p. 255, pi. 579. 
Sterna macrura, H. Saunders, P. Z. S. 1876, p. 650. 
S , October, sea-shore. 



1887.] MR. SMITH ON MOLLtJSCA FROM THE CAMEROONS. 127 

6. List of the Reptiles collected by Mr. H. H. Johnston on 

the Cameroons Mountain. By Gr. A. Boulenger. 

[Eeceived January 8, 1887.] 

1. Varanus niloticus (L,). 2000 feet. 
Hab. Whole of Africa south of the Atlas. 

2. Chameleon ovpenii, Gray. 2000 feet. 
Hab. Cameroon, Fernando Po, Gaboon. 

3. Urobelus gabontcus (A. Dum.). 2000 feet. 
Hab. Old Calabar to Gaboon. 

4. Naia haie (L.). 2000 feet. 

Hab. Whole of Africa south of the Atlas. 

5. Dendraspis angusticeps (Smith). 2000 feet. 

Hal. South Africa ; West Africa as far north as the mouth of the 
Niger. 

7. On the Mollusca collected at the Cameroons Mountain 

by Mr. H. H. Johnston. By Edgar A. Smith. 

[Eeceived January 12, 1887.] 

All tbe specimens collected by Mr. Johnston were from an altitude 
of from 7000 to 8000 feet. Of the seven species one only appears 
to be new. This is not surprising, as Dr. Buchholz made fine collec- 
tions in very much the same region some years ago, which were 
described by Dr. E. von Martens^. 

The species are : — 

1. Vaginula phuroprocta. Martens. 

2. Helicarion plicatulus, Martens. 

3. Helix {Trochonanina) percarinata, Martens. 

4. Stenogyra retifera, Martens. 

5. Stenogyra oleata, Martens. 

6. Streptostele buchholzi. Martens. 

7. Gibbus {U^deiitulina) johnstoni, sp. nov. 

The first two species were met with by Buchholz at Aburi ou 
the Gold Coast, the next four at Bonjongo in the Cameroons 
Mountain. 

The following is a description of the new species of Gibbus : — 

Gibbus (Edentulina) johnstoni. 

Testa anguste umbilicata, ovata, subtenuis, nitida, grisea; 

anfractus 6|^, celeriter crescentes, convexi, superiores regula- 

riter arcuatim et confertim striati, penult, et nltimus obsolete 

striati ; apertura inverse auriformis, albida, longit. totius f 

1 Monatsb. Akad. Wiss. Berlin, 1876, pp. 253-274. 



128 ON COIiEOPTERA. FROM THE CAMEROONS. [Feb. I, 

(squnns ; labrum tenue, angiiste reflexmn ; columella oblique 
contorta, expansa, umbilicum semiobtegetis. Longit. 30 m'dlim., 
diam. 16. 



Gibhus (Edentulina) johistoni. 

This is a smaller species than G. martensi. Smith, or G. insignis, 
Pfeiffer. It is also much more finely striated than the former and 
of a different shape, and has no plication on the columella as in the 
latter, which is an imperforate shell. 



8. On some Coleopterous Insects collected by Mr. H. H. 
Johnston on tlie Cameroous Mountain. By Charles 
O. Waterhouse. 

[Eeceived January 31, 1887.] 

The following Coleoptera were obtained by Mr. H. H. Johnston 
at altitudes from 8000 to 10,000 feet on the Cameroons Mountain : — 

1. ScARiTES ROTUNDicoLLis, Murtay. 

2. Two female examples of a Lamellicorn, apparently one of the 
Trichiidae ; 7 lines long, black, with deeply impressed strise on the 
elytra. It is very desirable to look for the male of this insect, which 
would doubtless be quite different in colour and perhaps in form. 

3. Batomene multispinis, Bates (Ent. Mo. Mag. xxi. 1884, 
p. 15), described from the Cameroons. 

4. Temnoscelis biemarginatus, Chevr. A single specimen 
in the British Museum from Old Calabar. 

5. A species of OtiorhgncJius closely resembling the European 
O. bisulcatus, but having the humeral angles slightly prominent, 
and the disk of the thorax with a short ridge. 

6. Epilachna, sp. 1 (In pieces.) 

From " Cameroons Mountain, 2000 feet, Oct. 1886 :"— 

7. Ceratorhina torquata, F. A common West-African 
species. 



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1887.] MR. F. DAY ON A HYBRID CLUPEOID FISH. 1^9 

9. On a supposed Hybrid between the Pilcliard [Clupea 
pilchardus) and the Herring^ (C. harengus), and on a 
specimen of Salmo purpuratas. By F. Day, C.I.E. 

[Received February 1, 1887.] 
(Plate XV.) 

In the mouth of September, 18S6, I received from Mr. Dunn, of 
Mevagisse)', in Cornwall, a hybrid Pilchard, and in December a 
second ; also the information that he had seen several, but that the 
scales had been injured in the remainder. The great interest in these 
fishes is that, although to a great extent the head most nearly 
resembles the Pilchard, the scales on the sides of the body show most 
remarkable differences — in the example which I propose describing 
consisting of 32 rows along the body and 8 rows in depth on the right 
side, whereas those along the left side are 51 in number and 10 rows 
in depth. 

Hybrids among the British Clupeidie have been observed ; thus the 
Alosa squamopinnata of Couch has been considered by Dr. Giinther 
to be a cross between the Pilchard and one of the Shads. 

.A cross between a Pilcliard and a Herring would apparently be 
rather remarkable, as Mr. Dunn found the eggs of the former floating ; 
however, I was informed by Professor Steindachner that those of the 
Sardine sink ; and as these are generally accepted as varieties of one 
species, it still seems doubtful whether the eggs float or sink. While, 
looking at the form of the head, it may be that the male Pilchard 
element had been prepotent in both fishes. 

The specimen figured (Plate XV.) has the following characters, and 
closely agrees with the second example, which is slightly longer, except 
that the size of the scales is reversed on the two sides of the body, being 
largest on the left side in the latter. 

D. 17. P. 15. V.8. C.23. L. 1. right side 32 ; 
left side 51. L. tr. right side 8 ; left side 10. 

inches. 

Entire length S'S 

Length of head I "7 

Length of caudal fin 1'5 

Length of pectoral fin 1 "0 

Height of body 1'7 

Eyes : diameter 0-4 of an inch, 0-5 inch from end of snout, and 
0-35 inch apart. The head is very similar to that of the Pilchard on 
the right side, while on the left the raised ridges on the opercles, 
although very distinct, are not so well marked as upou the opposite 
side. The form of the body is that of the Pilcliard. Gill-rakers : 
61 in the lower branch of the outer branchial arch, the longest 
being 0-35 inch, or not so long as the orbit. In a Herring the 
number of gill-rakers in the same position was found to be 48, and 

Proc. Zool. Soc— 1887, No. IX. 9 



130 MR. W. L. SCLATER ON [Feb. 1, 

two thirds as long as the eye ; in a Sprat 3.5, also two thirds as long 
as the orbit ; and in a Pilchard 71, longer than the eye. Thus these 
appendages in number in the hybrid (61) were less than in the 
Pilchard (71), but more than in the Herring(48) or in the Sprat (35) ; 
while their length did not quite equal tliat of a normal Pilchard. 
As to the character of these gill-rakers in the Pilchard, Sprat, and 
Herring : in the first the lateral denticulations are very minute, a little 
larger in the Sprat, and largest in tlie Herring ; to which last those 
in the hybrid had the greatest affinity. Fins. — In the hybrid the 
distance from the end of the snout to the commencement of the 
dorsal fin was 3*4 inches, the entire extent of the base of the fin being 
inserted midway between the snout and the root of the caudal fin ; 
lower lobe of the caudal the longer ; anal rays almost hidden by the 
scales. Scales. — The number of scutes 22 before and 14 behind the 
base of the ventral fin ; they are weak. As regards the scale3, two 
large rows exist just behind the head on the left side, and the 
remainder resemble to a great extent those of the Herring, but with 
the semicircular striae of the Pilchard. The scales on the right side 
are similar to those normally seen in the Pilchard. The number of 
scales along the body in Pilchards is from 29 to 30, in about 8 
vertical rows, 17-18 scutes before the ventral fin and 14 behind it; 
in the Sprat 47 scales along the body in 13 rows, 21-23 strong scutes 
before the ventral fin and 11-12 behind it; in the Herring the 
numbers of rows of scales along the body vary from .53-60, and there 
are 13 scutes behind the ventral fin. Colours. — On the left side was 
seen the beautiful purplish-golden hues of the Herring, but on the 
right side were the silvery colours of the Pilchard. 

Salmo purpuratus. 

This specimen, which is 8*5 inches long, died at South Kensington 
in August 1885. It was one of the fishes raised from the eggs brought 
over from Canada by Mr. Wilmot in 1883, which were described in 
the Society's 'Proceedings' for 1884, p. 24. Originally imported 
as supposed eggs of the Salmon, the edge of the adipose dorsal fin 
in the fry showed the orange tints of a Trout, while the par-bands 
were from 7 to 10 and averaged 85. It is interesting, because spe- 
cimens have been turned into the Thames, and were asserted to be 
Land-locked Salmon, which is an error of identification. 



10. Notes on the Peripatus of British Guiana. 
By W. L. Sclater, B.A., F.Z.S. 

[Eeceived January 31, 1887.] 

During my recent stay in Demerara I was fortunate enough to 
procure a considerable number of specimens of a species of Peripatus. 
This singular form was first discovered in British Guiana by Mr. im 
Thurn, who sent examples home to Prof. Moseley. But the bottle 
containing the specimens was broken before arriving in England, and 



1887.] PERIPATUS FROM BRITISH GUIANA. 131 

the contents were dried up. Of the Peripati which I obtained twenty 
individuals were brought to England alive, but were unfortunately 
found to be much affected by the cold, and were tberefore killed and 
preserved immediately on arrival. I also brought with me four 
other specimens that had been preserved in British Guiana. 

All the specimens which I obtained were females, and all of them 
contained embryos. A\\ the specimens examined, both large and 
small, including those taken from the uterus, were found to have 30 
pairs of legs, and of course a pair of oral papillae. In this respect 
they differ from the form of Caraccas described by Ernst, in which, as 
he states, the young ones have only 29 pairs of legs, while the adult 
specimens possess 31 pairs. The colour of the Demeraran Peripatus 
is a dark brick-red above and pinkish below with a dark suffused 
median line on the dorsal surface, such as Ernst (25) described in his 
specimens. The antennae are very much darker than the rest of the 
body, in fact they are quite black. The body, as in all other forms of 
Peripatus, is divided into numerous rings by lines of small warts, 
about 10 to 12 rings going to each segment ; the legs and antennae 
are also ringed, and the former bear the usual pair of hooks. 

In the living animal the colour is intermediate between the colour 
of the two specimens now exhibited — that preserved in spirit being 
of a darker, and that preserved in Pereney ' fluid being of a lighter 
hue, than that of the living animal. The adult specimens vary from 
2-25 to about 2*50 inch in length. It is useless to give exact 
measurements, since not only do the animals contract when preserved 
in spirit, but even the living animals vary greatly in size at 
different times. 

The question as to what species the Demeraran Peripatus should 
be referred is by no means an easy one. Specimens of Peripatus have 
been obtained from the following places in the West Indies and South 
and Central America: — 

(I) St. Vincent's, W. I. ; with 33 pairs of legs. Guilding (1). 
(2J Cayenne ; with 29 pairs of legs. Audouin and Milne-Edwards 

(2) 

(3) Lake of Valencia, Venezuela. Wiegmaun (4). 

(4) Chili. Gay (12). 

(,.5) St. Thomas, W. I. Moritz (5). 

(6) Colonia de Tovar, Venezuela. Grnbe (11). 

(7) Santarem, Amazons ; 3 1 pairs of legs. Moseley (22). 

(8) Nicaragua. Belt (17). 

(9) Caraccas, Venezuela. Ernst (25). 

( 10) Trinidad (2 species). Kennel (30) and (31). 

(II) Island of Marajo, Amazons. Branner (34). 

(12) Dominica. Bell (28). 

(13) Porto Rico. Peters (23). 

P. torquatus, a species discovered by Kennel (31) in Trinidad, 
and described by him, is easily distinguished by its large size 
(15 mm.), the number of its pairs of legs (41-42), and by its yellow 

' Pereney fluid is a hardening fluid composed of chromic acid '5 p. c. sol., 3 
parts ; nitric acid 10 p. c. sol., 4 parts ; spirit 90 p. c, 3 parts. 

9* 



132 MR. W. L. SCLATER ON [Feb. 1, 

collar. P. blainvUlii, the species described by Gay from Chili, 
seems also to be distinct, as posessing only 1 9 pairs of legs. 

All the other forms from the above-mentioned localities, including 
Kennel's second species from Trinidad, seem, so far as one can judge 
from the descriptions, to resemble one another very closely, except 
as regards slight variations in the number of pairs of legs. Thus 
Guildiiig's species {P . juliformis) is described as possessing 33 pairs 
of legs ; vphile the form from Caraccas is said by Dr. Ernst to have 
31 pairs in the adult, and only 29 when first born. All my specimens 
from Demerara of all ages agree in having 30 pairs of legs. 

Another point in which the Demeraran form seems to differ from 
the other forms described is that the colour of the antennae is black. 
This point is not specially mentioned in the descriptions of the other 
American Peripati. 

I have also examined the examples of Peripatus in the British 
Museum. Of all the examples of the genus in the National collection 
there is only one specimen which seems to resemble my form ; it is 
that labelled " Peripatus from Dominica, found under logs." The 
animal in question was obtained in Dominica and presented to the 
British Museum by the late Mr. G. F. Angas, C.M.Z.S., and has 
been noticed by Prof. F. Jeffrey Bell (28). 

The Peripatus from Dominica resembles the Demeraran form in 
the following points : — the black antennae ; the general colour, so 
far as can be judged from the spirit-preserved specimens ; the number 
of legs (30 pairs) ; and also in another point which I have not 
hitherto mentioned, but which seems to otfer characters useful for 
distinguishing the various species : this is the shape of the slits on 
the under surface of the feet. 

In all the American specimens examined by me at the British 
Museum this slit is split-shaped ; but in my specimens and in 
that from Dominica the openings are in many cases rounded, and 
sometimes have attached to them a bladder-shaped appendage, as 
mentioned by Prof. Bell (28). 

It seems to me therefore that there are only three species oi Peri- 
patus yet satisfactorily determined in South America. 

1. P. torquatus, Kennel, from Trinidad. 

2. P. blainvillii. Gay, from Chili. 

3. P. edwardsi, Blanchard {juliformis, Guilding ?), from Cay- 
enne, British Guiana, Venezuela, Nicaragua, and several of the 
West-India Islands. 

To these three species must be added a fourth, from Dominica and 
British Guiana, distinguished by the following points : — 

(1) The black antennae. 

(2) Thirty pairs of feet and one pair of oral papillae. 

(3) The darker and redder colour; the other forms being a dirty 
brown colour as far as can be seen in the spirit specimens. 

(4) The rounded openings to the foot-pits. 

(5) The black marking in the median dorsal line in these forms, 
■which is much more definite than in any of the others from South 
America, 



1887.] PERIPATUS FROM BRITISH GUIANA. 133 

I do not give a name to this Dominican and British-Guianan 
Peripatus, since 1 understand that Mr. Sedgwicii is about to 
pubhsh a monograph on the species of the genus Peripatus, and 
will include in his work a description of the specimen from Dominica 
in tlie British Museum. 

All the specimens of Peripatus obtained by me were found, with 
one exception, in the grounds round Mr. im Thuru's house, Mac- 
casseema, on the Pomeroon River. Maccasseema is situated on 
the top of a sand-hill about 30 feet above the river, and is sur- 
rounded on all sides by the swampy forest, except in front, where 
it faces the river. The specimens were all found under rotten 
logs of wood, or under the decaying stalks of the Cokerite Palm 
{Maximiliana martiana). I never saw one actually in the rotten 
wood, as has been described by some previous observers. 

The single exception was found about a mile from Maccasseema, 
up a creek running into the river Pomeroon. This individual was also 
found under a more or less rotten log close to an Indian house. 

Specimens of Peripatus were exceedingly scarce, and it took a 
long time to collect even the few I brought home. 

I should mention that examples of Peripatus have also been 
obtained in Demerara by Mr. Qnelch, the Curator of the Georgetown 
Museum, who found them about twenty miles from Georgetown on 
the Hoorubea Creek (36). 

In offering these preliminary notes on this most interesting animal, 
I have not entered into further details, because Prof. Moseley and 
Mr. Sedgwick are about to publish an account of the different 
species of Peripatus, and will incorporate their observations on the 
present form into their work. But before concluding I must express 
my thanks to Mr, im Thurn for all the help he gave me in my 
collecting, more especially for allowing me the use of Douglas, the 
captain of his Indian boat's crew, as collector, for to his sharp eyes 
I owe most of my specimens. 



APPENDIX. 



The numbers appended to the authors' names in this paper refer 
to the following list of publications, which forms, I believe, a nearly 
complete bibhography of original works on Peripatus. To most of 
the titles I have added a few remarks explaining the contents of the 
memoirs. The publications that I have not been able to examine at 
first hand are marked with an asterisk. 

(1) GuiLDiNG, L. Mollusca Caribbaeana; an account of a new 
genus of Mollusca. Zool. Journ. ii. pp. 443-444, pi. xiv» 
1826. 
Contains the original description of the genus Peripatus and 
species P. juVformis, found by the author in the forests of St. Vin- 
cent. The author considered it an aberrant form of slug. A fair 
coloured plate is given. 



134 MR. W. L. SCLATER ON [Feb. 1, 

(2) Atjdouin ct Milne-Edwards. Classification des Annelide?, 

etc. Ann. Sc. Nat. xxx. pp. 411-414, pi. xxii. 1833. 
The authors show that Peripatus must be placed among the 
"Annelides Errantes." The specimens described were obtained 
from the River Appronague, in Cayenne. 

(3) Gervais, p. Etudes pour servir a I'histoire naturelle des 

Myriapodes. Ann. Sc. Nat. (2) vii. pp. 35-60. 1837. 
The author believes Peripatus to be a transitional form between 
the Mjriapods and Chsetopods ; he also quotes a MS. description 
by Blainville of a second species (P. brevis) from the Cape of Good 
Hope. 

(4) WiEGMANN, A. F. A. Einige Bemerkungen iiber Guilding's 

Peiipatws. Arch. f. Nat. (Wiegmann), iii. pp. 195-200. 
1837. 
Description of certain specimens of Peripatus from the Lake of 
Valencia, in Venezuela. 

(5) MoRiTZ, C. Noch einige Worte iiber PmjofliM«, Guild. Arch. 

f. Nat. (Wiegmann), v. pp. 175-1 70. 1839. 
Remarks on the habits and life-conditions of specimens of Peri- 
patus found in St. Thomas. 

(6) De Blainville. Dictionnaire des Sciences Naturelles. Sup- 

plement, tom. i. p. 237. Paris, 1840. 
In his article " Animal " de Blainville institutes a special division, 
" Les 3JaIacopodes" of his Type ii. "Entomozoaires," for the genus 
Peripatus. 

(7) Milne-Edwards, H. Note sur le Peripate juliforme. Ann. 

Sc. Nat. (2) xviii. pp. 126-128. 1842. 
The author reiterates his opinion of the Annelidaa nature of 
Peripatus, 

(8) Blanchard, E. Recherches sur Torganisation des Vers. 

Ann. Sci. Nat. (3) viii. pp. 119-149. 1847. 
On pp. 137—141 is given the history of the four species known at 
that time — P. juli/ormis, edwardsi, blainvillii, and brevis. The 
form is considered by the author to be related to Annelids. 

(9) Qviatrefages, A. DE. Memoire sur la famille des IlermeUiens. 

Ann. Sci. Nat. (3) x. pp. 5-58. 1848. 
Quatrefages (p. 5G) considers Peripatus to be a worm of aberrant 
form, dittinctly related to this group (e. g. Hermellea). 

(10) Milne-Edwards, Quatrefages, et Blanchard. Recher- 
ches anatomiques et zoologiques fait pendant un voyage sur la 
cote de Sicile, part iii. p. 61, pi. i. fig. 2. Paris, 1849. 

Blanchard forms a new species (P. edwardsi) for the reception 
of Milne-Edwards's form from Cayenne ; he also mentions Gay's 
species, P. blainvillii, afterwards described (12). and gives some 
account of its anatomy. 



1B87.] PERIPATUS FROM BRITISH GUIANA. l.'<5 

(11) Grube, E. Untersuchungen iiber den Bau von Peiipatus 
edwardsii. Miiller's Arch. Auat. Phys. 1853, pp. 322-360, 
Taf. ix., X. 1853. 

A description of the anatomy of Peripatua from specimens obtained 
at Colonia de Tovar, in Venezuela. 

(12) Gay, C. Historia fisica y politica de Chile. Fauna, Vol. iii. 
Atlas, Annelides, Lam. iii. fig 2. 1854. 

On page 58 is a description of a new species {Peripatus blainvillii), 
with 19 pairs of legs, from Chile. In the Atlas, it may be observed, 
this species is figured with varying numbers of pairs of legs, in one 
case 30 and another 29. 

(13) QuATREFAGES, A. DE. Histoirc Naturelle des Anneles. 
Tome ii. Paris, 1865. 8vo. 

On page 6/5 is an account of the genus Peripatus and of the 
species then known. 

(14) Grube, E. Reise derosterreichischenFregatte Novara. Zool. 
Theil ii. Anneliden. Wien, 18G7. 

Description of P. capensis found near Coustautia at the Cape. 
See p. 4, pi. iv. figs. 3, 4 a. 

(15)* Sanger. Description of a Peripu^Ms from Australia. Trans- 
actions of the Russian Assembly of Naturalists, held at 
Moscow in 1867. Moscow, 1869. 
Description of P. feuckarti from Australia, and of the anatomy of 
P. capensis, in Russian. 

(16) Leuckart. Bericht iib. Leist in d. Naturgeschichte der 
Niederen Thiere wiihrend der J. 1868-1809. Arch. f. Nat. 
(Troschel), xxxv. pt. 2, pp. 277-278. 1869. 

A resume of Sauger's paper, containing an account of the anatomy 
of Peripatus capensis and of the new species Irom Australia, Peri- 
patus leuckarti. 

(17) Belt, T. The Naturalist in Nicaragua. 8vo. London, 
1874. 

Mention is made (p. 140) of Peripatus in Nicaragua, though not 
by name. The specimen procured by Belt was afterwards identified 
by Prof. Moseley (22). 

(18) Moseley, H. N. On the Structure and Development of 
Peripatus capensis. Phil. Trans, clxiv. pis. Ixxii.-lxxv. 
pp. 757-782 ; and Proc. R. S. xxii. pp. 344-350. 1874. 

General account of the anatomy and development of the Cape 
species of Peripatus from specimens obtained during the ' Chal- 
lenger' Expedition. 

(19) Button. F. W. Oa Peripatus novcB-zealandice. Ann. Mag. 
N. H. (4) xviii. pi. xvii. pp. 361-369, 1876 ; also op. cit. 
(4) XX. pp. 81-83, 1877; and op. cit. (5) i. pp. 204-206, 
1878. 

Description of the New-Zealand species of Peripatus, with au 



136 MR. w. L. scLATER ON [Feb. 1, 

account of its habits and anatomy, and a few words on its develop- 
ment. 

(20) MosF.LEY, H. N. Remarks on Observations by Capt. Hutton, 
Director of the Otago Museum, on Peripatus novoe-zealandice, 
with notes on the Structure of the Species. Ann. Mag. N. H. 
(4)xix. pp. 85-91. 1877. 

A criticism of Hutton (19\ with additional remarks on several 
jioints in the anatomy of P. novce-zealandice not mentioned by him. 

(21) Balfour, F. M. On certain Points in the Anatomy of Peri- 
patus capensis Proc. Cambr. Phil. Soc. iii. pp. 266-269; 
Quart. J. Micr. Sc. xix. pp. 431-433. 1879. 

Description of the renal segmental organs and of certain points in 
the anatomy of the nervous, system of Peripatus. 

(22) MosELEY, H. N. Notes on the Species of Peripatus, and 
especially on those of Cayenne and the West Indies. Ann. 
Mag. N. H. (.5) iii. pp. 263-267. 1879. 

Contains a history of the genus and a discussion as to the number 
of species in South America ; also notes on two specimens, one from 
Santarem, on the Amazons ; the other from Nicaragua, collected by 
Mr. Thomas Beh(I7). 

(23) Peters, W. Ueber die Arten von Peripatus. SB. nat. Fr, 
Berlin, 1880, pp. 28-29. 1880. 

A short account of the species then known (four), and remarks 
on the variation of the number of pairs of legs. Records tiie exist- 
ence of specimens in the Berlin Museum from Porto Rico, Surinam, 
and Venezuela, 

(24) Peters, W. Die Variation der Fusszahl bei Per?)ja^M5 ca/3e««is, 
Giiibe. SB. nat. Fr. Berhn, 1880, pp. 165-166. 1880. 

Records the variation in the number of pairs of legs in a series of 
Peripatus from the Cape of from 22 to 17 pairs. 

(25) Ernst, A. Some remarks on Peripatus edwardsii, Blanch. 
Nature, xxiii. pp. 446-448. 1881. 

An account of specimens found at Caraccas, in Venezuela. 

(26) MosELEY, H. N., and Sedgwick, A. Note on a discovery, 
as yet unpublished, by the late Professor F. M. Balfour, con- 
cerning the existence of a Blastopore, and on the Origin of the 
Mesoblast in the Embryo of Peripatus cajjeiisis. Proc. R. 
S. xxxiv. pp. 390-393. 1882. 

(27) Balfour, F. M. The Anatomy and Development of Peri- 
patus capensis ; edited by Prof. H.N. Moseley and A. Sedg- 
wick. Quart. J. Micr. Sc. xxiii. pp. 213-259, pis. xiii.-xx. 
1883. 

Descrijition of the anatomy and some account of the developn^ent, 
with a coloured plate, of P. cape7isis. 

(28) Bell, F. J. Note on a Peripatus from the Island of Domi- 
nica, West Indies. Ann. Mag. N, II. (5) xi. [>. 388. 1883. 



1887.] PERIPATUS FROM BRITISH GUIANA. 137 

(29) Gaffron, E. Beitrage zur Anatomie und Histologie von 
Peripatus. Zool.Beitr. (Schneider), i. Taf. vii.-xii. pp. 33-60. 
1883. Also ^?H. ci<. Taf. xi., xii., xiii., pp. 14.')-162. 1885. 

Account of the anatomy and more particularly the histology of 
Peripatus edwardsii from Trinidad, with 32 pairs of legs. 

(30) Kennel, J. Entwicklungsgeschichte von Peripatus. Zool. 
Anz. vii. pp. 531-537. 1883. 

A preliminary notice, containing a description of P. torquatus 
from Trinidad. 

(31) Kennel, J. 'Entmcklnngsgeschichte \on Peripatus edwardsii, 
Blanch., \mA Periptaus torquatus, n. sp. Theil I. Mit Taf. v. 
bis xi. Arbeit, zool.-zoot. Inst. Wiirzburg, vii. pp. 95-228. 
1885. Theil II. Mit Taf. i. bis vi. Arbeit, zool.-zoot. Inst. 
Wiirzburg, viii. pp. 1-93. 188G. 

An account of the development of the American species of Peri- 
patus, which are characterized by the absence of food-yolk in the 
ova, and by the presence of a (so-called) placenta. The specimens 
examined were obtained from Trinidad. 

(32) Sedgwick, A. On the Fertilized Ovum and Formation of 
the Layers of tlie South-African Peripatus. Proc. Roy. Soc. 
xxxix. pp. 239-244. 1885. 

Preliminary account of no. (33). 

(33) Sedgwick, A. The Development of the Cape Species of 
Peripatus. Part I., with pis. xxxi., xxxii. Quart. J. Micr. 
Sc. XXV. pp. 449-446. 1885. Part II., with pis. xii.-xiv. 
Quart. J. Micr. Sc. xxvi. pp. 175-212. 1886. 

The first part contains the first mention of P. ba/fouri, distin- 
guished by having 18 pairs of legs ; it also contains an account of 
the generative organs, segmentation, and general development of the 
embryo. Part II. contains a further account of segmentation and 
early stages. 

(34) Branner, J. C. Peripatus in the Island of Marajo, Ama- 
zons. Nature, xxxiv. p. 496. 1886. 

(35) HoRST, R. On a specimen of Peripatus, Guild., from Su- 
matra. Notes Leyd. Mus. viii. pp. 37-41, pi. ii. figs. 1-5. 
1886. 

Descrijition of a specimen of Peripatus with 24 pairs of claw- 
bearing legs, from Sumatra. 

(36) QuELCH, J. J. Peripatus in Demerara. Nature, xxxiv. 
p. 288. 1886. 

(37) Stuhlmann, F. Die Reifung des Arthropodeneis. Pp. 1- 
128, Taf. i.-iv. Freiburg-i.-B. 1886. 8vo. 

Account of the ovary and ovarian ovum of P. edwardsii. Pp. 89- 
93. 



138 THE SECRETARY ON ADDITIONS TO THE MENAGERIE. [Feb. 15, 

February 15, 1887. 
Prof. W. H. Flower, LL.D., F.R.S., President, in the Chair. 

The Secretary read the following report on the additions to the 
Society's Menagerie during the month of January 1887: — 

The total number of registered additions to the Society's Mena- 
gerie during the month of January was 38. Of these 6 were by 
birth, 21 by presentation, 5 by purchase, I by exchange, and 5 were 
received on deposit. The total number of departures during the 
same period, by death and removals, was 119. 

The most noticeable additions during the month were : — 

1. Two Blakiston's Owls {Bubo blakistoni) from Japan, presented 
by J. H. Leech, Esq., F.Z.S., January 20th, new to the Society's 
Collection. 

Mr. Leech informs me that he procured the two specimens in 
question from Mr. H. Henson of Hakodate, Yesso. Mr. Ilenson had 
bought them from a native hunter, who took them for young eagles, 
wliich are common in Ytsso. Their exact locality is the lakes 20 
miles north of Hakodate, and this is a new locality for the species, 
of which, Mr. Leech believes, only four specimens were previously 
known ^ 

Mr. J. H. Gurney, F.Z.S., has kindly furnished me with the 
ftllowing notes upon these birds: — 

" Blakiston's Eagle-Owl appears to belong to the Bubonine genus 
(or subgenus) Pseudoptynx of Kaup, instituted by that author for 
the reception of P. phi/ippensis (Gray), and readily distinguislied 
from Bubo in having the toes bare, although the tarsi are feathered. 

" P. philijJijensis is a native of the island of Luzon, from which I 
have seen specimens, and was figured by the late Lord Tweeddale in 
the Society's 'Transactions,' vol. ix. pi. xxv. fig. 2. 

"The only other sfiecies of the genus is P.gurneyi, Tweed., from 
the island of Mindanao, which was figured by Lord Tweeddale in the 
'Proceedings' of the Zoological Society, 18/8, pi. Iviii. 

" The localities inhabited by the three species of the genus Pseu- 
doptynx appear to indicate that they form a natural group, geogra- 
phically as well as structurally."— j! H. G., Feb. 26, 1887. 

2. Three Hooker's Sea-lions {Otaria hookeri), presented by the 
Hon. W. J. M. Larnach, C.M.G., Minister of Marine of New 
Zealand, received the 2Cth January. 

Sir F. D. Bell, the Agent-General for New Zealand, informs me 
that these animals, which were captured at the Auckland Islands by 
Capt. John Fairchild, Master of the New-Zealand Government 
steamer ' Hinemoa,' were originally four in number (two males and 
two females), but that one died on the voyage. 

It is very difficult to settle the species of Otaria without reference 
to the form of their palates and dentition ; but, judging from the 

' Cf. Seebohm, ' Ibis,' 1884, pp. 42, 183, pi. vi. ; id. P. Z. S. 1883, p. 406. 



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Miiil.<ii-u BfiMt, imp. 



ech:nodepims of the Andaman islands. 



1887.] ON ECHINODERMATA FROM THE ANDAMAN ISLANDS, 139 

locality and their appearance, these Sea-lions must be referred to 
the species of the Auckland Islands, upon which Mr. J. W. Clarke, 
F.Z.S., made his valuable communication in 1873 (see P. Z. S. 1873, 
p. 750), and should be called Otaria hookeri. 

The largest male is nearly equal in bulk to our old male O.jubata, 
but has much shorter front flippers and rather longer external ears. 

3. A Blue Penguin {Eudyptula minor), from Cook's Straits, New 
Zealand, presented by Mr. Bernard Lawson, January 26th, being 
the first example of this interesting little Penguin that has been 
received by the Society. 

The following papers were read : — 

1. Keport on a Collection of Echinoderraata from the 
Andaman Islands. By F. Jeffrey Bell_, M.A., Sec. 
R.M.S., Professor of Comparative Anatomy and Zoology 
in King's College, London. 

[Eeceived January 18, 1887.] 
(Plate XVI.) 

Dr. John Anderson, F.R.S., Superintendent of the Indian Museum, 
Calcutta, was lately kind enough to excite the interest of Col. Cadell, 
V.C., in the marine zoology of the Andaman Islands, which are at 
present under his charge, and to present to the British Museum the 
collections thus made. The following contains a report on the Eclii- 
noderms, which Dr. Anderson has asked me to draw up. 

The condition in which the specimens reached England reflects great 
credit on Mr. Booley, who made the collections for Dr. Anderson. 

There are in all fifty species of Echinoderms, of which no less 
than twenty-two are Holothurians ; the bulk of what follows will 
treat chiefly of these interesting but difficult forms, which are abun- 
dantly found in tlie Eastern seas. Of the Asteroids, Linclia Icevi- 
gata was exceedingly abundant, there being twenty exainples of 
it, and one only of L. pacifca ; of these twenty examples, one 
was four-rayed. Scy taster nova-caledonice was not rare ; Culcita 
was represented by handsome species. Of two of the most difficult 
genera, Linckia, Astropecten, there is in each case a single example 
of a form unknown to me ; 1 cannot associate either with a described 
congeneric form, but, on the other hand, I am not satisfied that they 
are the representatives of " new species." 

Among Ophiuroids, the only noteworthy point is the complete 
absence of Ophiothrix from the present collection. There is but 
one Crinoid. 

It is to be regretted that it is impossible for me to compare the 
results of a collection at Mergui with that now before me, my report 
on the Holothurians collected by Dr. Anderson being as yet the only 
portion of the account of Echinoderms which has appeared in the 



1-10 PROF. F. J. BELL ON ECHINODERMATA [Feb 15, 

Journal of the Linnean Society i; with the contents of this paper 
the reports of Prof. Duncan, Dr. H. Carpenter, and Mr. Sladen on 
the other groups of Mergui Echiuoderms should, on their publica- 
tion, be consulted. 

I have tried to arrange the information to be given in the shortest 
and clearest way possible, giving first a list of the species, then 
remarks on those to which it is important to direct special attention. 

I. Crinoidea. 

1. Antedon, sp. 



II. ASTEROIDEA. 



2. Acanthaster ecbinites, E. ^- 8. 

3. Fromia iudica, Perrkr. 

4. tuniida. Bell. 

5. Linckia Irevigata, L. 

6. paciiica, Gray. 

7. Scytaster novss-caledoiii£e, 

Perrier. 



8. Culeita grex, M. Tr. 

9. schmideliana, Betz? 

10. sp. (" Eandasia granulata.") 

11. Astropecten polyacanthus, il/. Tr. 

12. sp. 

13. Archaster typicus, M. Tr. 



III. Ophiuroidea. 



14. Pectitmra gorgonia, M. Tr. 

15. Opbiolepis aiimilosa, M. Tr. 

16. Ophiocoma scolopendrina, Lamk, 



17. Ophiocoma tetbiops, Lutlcen. 

18. Opbiomasiix annulosa, M. Tr. 

19. Opbiaracbna inerassata, ^I. Tr. 



IV. ECHINOIDEA. 



20. Pbyllacantbus verticillatus, Lmk. 

21. Diadema setosuin, Gray. 

22. Astro]3yga radiata, Lcske. 

23. Toxopueustes pileohis, Lamk. 

24. Ecbinometra lucmiter, Lcske. 



25. Colobocentrotus atratus, L. 

26. Echinoneus cylo.stoimis, Lcske. 

27. Metalia steriialis, Lamk. 

28. Moira stygia {Liitken, MSS.), 

A.Ag. 



V. HOLOTHURIOIDEA. 



38. Holotburia lineata, Lvdwig. 

39. maculata, Branclf. 

40. mariuorata, Jmjer. 

41. ruoiiacaria, Lcssoii. 

42. papillata. 

43. Tagabunda, Sclcnka. 

44. Sticbo]nis cbloronotus, Brandt. 

45. variegatus, Semper. 



29. Chirodota rufescens, Brandt. 

30. Haplodactjla aiidamaiiensis. 

31. Pseudocucuniis acicula, iicmper. 

32. Muelleria luaiiritiana, Q. ^' G. 

33. miliaris, Q. ij' G. 

34. Holotburia albida. 

35. atra, Jdger. 

36. cadelli. 

37. impatiens, Forskal. 

In addition, there are five specimens belonging to as many species 
o{ Holothuria as to which I must reserve an ojiinion till I have, as 
I hope to have soon, a larger series before me. 

Crinoidea. 
Antedon, sp. 

This Antedon is represented also in the collection made at Mergui 

^ J. L. S. xxi. p. 25 ; tbe wbole of vol. xsi. of that Journal will be devoted 
to tbe fauua of Mergui. 

^ Prof. Loveii assures me that I was quite wrong in ascribing to Bruzelius 
the tract that bears bis name (Ann. N. H. (b) ix. p. 166). 



188/.] FROM THE ANDAMAN ISLANDS. 141 

by Dr. Anderson; and Dr. Herbert Carpenter, F.R.S., has promised 
to discnss its relations to A. palmata in the report on the Crinoids 
of Mergui which he has in preparation. 

ASTEROIDEA. 
ACANTHASTER ECIIINITES. 

M. de Loriol has lately pointed out ' that the species oi Acanthaster 
fonnd at tlie island of Mauritius is not, as has been supposed, 
A. echinites ; a comparison of liis description and Mauritian speci- 
mens with the figures of Ellis and Solander and examples from the 
Andamans will be sufficient to show the student the distinctness of 
the species. 

As the difference has only lately been pointed out, and the con- 
fusion cleared up, it is as yet too early to say whether A. echinites 
belongs to the eastern, and A. mauritiensis to the western side of 
the Indian Ocean, or whether their areas of distribution overlap. 

Fromia indica. 

I have elsewhere '^ given my reasons for regarding this species, 
described by Piof. Perrier as six-rayed, as being normally quinque- 
r.ndiate ; a five-rayed specimen in the present collection has R equal 
to 33, and ?=9. 

With it are two specimens which possibly belong to a different 
species of the same genus ; they are smaller and are still quite 
spiny. 

Culcita schmideliana. 

There is a very remarkable specimen which I fancy I am hardly 
wrong in describing in detail ; another is of the more ordinary 
character. 

Almost round ; the apices of the ambulacra just touch the 
equator, so that R is almost exactly equal to r ; the ambulacra 
narrow rapidly after reaching the actinal periphery. The ordinary 
arrangement of the adambulacral spines is as follows : — In the inner- 
most row four subequal spines, beside which there may be a fifth 
smaller ; outside of and touching these there may be one large or 
two smaller spines, and either one or both occupy as much of the 
side of the groove as do the four spines internal to them ; outside 
of the second there is a third row which is more irregular, especially 
in the region of the actinostome. All the spines are stout, and 
more or less rounded at the tip. The interambulacral area, which is 
thickly covered with flat-headed grains, is almost perfectly triangular 
in shape ; the number of grains in a patch varies ; the patches are 
more closely packed in the n)iddle than at the sides of the interambu- 
lacral triangle, and scattered among them are the ordinary granules. 
Peripherally the patches of grains cease somewhat rapidly ; a band, 
bare of patches, but granular and with sparsely scattered tubercles, 

^ M6m. Soc. Phys. Geneve, xxix. no. 4, p. 6. 
= Proc. Zool. Soc. 1882, p. 12.3. 



142 PROF. F. J. BELL ON ECHINODERMATA [Feb. 15, 

separates them from the poriferous area ; this last extends down to 
the edge of the actinal surface. 

On the abactinal surface there are scattered tubercles and large 
poriferous patches ; over the whole there is a uniform granulation ; 
no pedicellarise were detected. 

The madreporic tubercle is large and prominent. 

Colour, in spirit, light yellow, the poriferous patches darker. 

Measurement round the equator 560 mm. ; height 75 mm. along 
the longest axis. 

The most interesting points with regard to this species are such 
as bear on its relation to the Echinoidea, Those who accept the old 
doctrine of Cuvier and Duvernoy, restored in these days by Prof. 
Haeckel, which explains the constitution of the Echinoderm by the 
hypothesis that it consists of several fused persons, have found in 
Culcita the form that seemed to show how the free arms of the 
Starfish might pass into the compact form of the Urchin. This theory 
of Echinoderm constitution does not recommend itself to me ; and 
tlie present species seems to justify the hesitation which one feels in 
accepting it. 

While in flattened or irregular Echinoids there is a tendency for 
the ambulacra to shorten towards the aboral pole, here the tendency 
is for the shortening to be towards the mouth. In other words, the 
most extreme Asterid which we know, though it has a remarkable 
general resemblance to an Echinoid, is, in its essential morphological 
points, further from it than is a typical Asterid. 

Culcita grex. 

"With a little hesitation I refer a single specimen to this species ; 
the variability of the forms of the genus Culcita is obviously very 
great, and a careful revision of the species with the aid of a large 
number of specimens is a pressing necessity. The collection in the 
National Museum is not yet sufficiently large to justify me in under- 
taking the task. 

Culcita, sp. 

There are two specimens of what would, a short time ago, have 
been set down as Randasia granulata. As, however, M. de Loriol 
has lately shown, the form so called by the late Dr. Gray is really 
a young stage of Culcita. It will be remembered that Prof Perrier 
has expressed himself in a similar sense. Further series are required 
before the several stages of each species can be accurately defined. 

Echinoidea. 

DiADEMA SETOSUM. 

I am really very doubtful as to the specific identity of two small 
specimens, the spines of which are banded light and dark, and present 
the charactersof Dr. Gray's "C'a/wrtHMSflwne//«/a," with the adult large 
black-spined forms which are assumed by D. setosum. However, I 
have not sufficient evidence to justify me in attempting to refute the 



18S7.] FROM THE ANDAMAN ISLANDS. 143 

conclusions formulated by Mr. Alex. Agassiz in his ' synonymy ' 
of this species. The larger specimens collected are magnificent 
examples of this interesting species. 

MoiRA STYGiA. (Plate XVI. figs. 1-3.) 

Being in some doubt as to whether I had before me the species 
described by Dr. Liitken, I sent the drawings here given to that 
accomplished zoologist with the request that he would compare them 
with the type in the Copenhagen Museum. Dr. Liitken writes : — • 
" The specimen from Zanzibar is much smaller than your figure 
[which bore the mark x U] — 24 mm. in length and 14 mm. in 
height. The anterior lateral ambulacra are more bent, the posterior 
shorter than in your figure ; the posterior excavation of the shell 
below the periproct less crested than in your figure. But these 
differences might be those of age." The most striking difference 
to which this obliging communication directs attention is the great 
difference in the proportion of height to breadth ; however, in a 
specimen from an unknown locality, which I am inclined to place 
also in this species, the proportion of height to length is about 
the same, for it is as 37 to 30, whereas in the Andaman species it 
is as 33-5 to 21 ; and tiie very same is true also of the specimen 
whose dimensions are given in the ' Revision of the Echini,' where 
the height is to the Ion"; diameter in the ratio of 40 to 49'5. 

The other dimensions of the Andaman specimen are : — Breadth 
28"5 ; length of antero-lateral ambulacrum 13; of the postero- 
lateral ambulacrum 10 ; length of anal area 4 ; breadth of do. 3 
millim. 

It is now for the first time figured, and is the first specimen of 
the genus that has been shown with the spines on. 

The discovery of this species at the Andamans extends its range, 
though not indeed in any unexpected way ; hitherto specimens have 
been known only from the Red Sea and Zanzibar. 

HOLOTHTIRIOIDEA. 

Haplodactyla andamanensis. (Plate XVI. fig. 4.) 

Body elongated, tapering posteriorly ; the skin of a slightly reddish- 
grey hue, darker above than below. 

Tentacles ? (retracted). 

The body-wall is thin ; the ossicles of the oesophageal ring are 
elongated, rather stout, deeply grooved on the outer surface ; the 
radials are longer than the interradials, and have a bifurcated distal 
tail. One Polian vesicle. Genital tubes numerous, long, well 
developed, purple in colour. Lungs extend to anterior end of body, 
two well but unequally, and one poorly developed lung-trunk ; the 
last does not exteid far forwards. Biscuit-shaped or dumbbell-like 
spicules (Plate XVL fig. 4). 

By the forms of its spicules it is distinguished easily from //. mol- 
padoides, and by them and the tapering of the hinder end from H. 
australin. 



144 PROF. F. J. BELL ON liCHINODERMATA [Feb. 15, 

PsEUDOcucuMis ACicuLA. (Plate XVI. fig. 5.) 

A single specimen of this rather rare species, the spicules of which 
I have had figured, as their representation seems to be more satis- 
factory than those of Prof. Semper. 

MUELLERIA MILIARIS. 

Some of the specimens which I associate under this name differ 
from M. lecanora in that the region of the anus is not lighter than 
the rest of the dorsal surface ; on the other hand, the sharp distinc- 
tion between the dark brown of the bivial and the light colour of the 
trivial surface is an indication of affinity to M. lecanora. Consider- 
ing the closeness of the resemblances and the slightness of the 
differences between M. miliaris and M. lecanora, I feel inclined to 
suggest that the species should be united. One specimen is of a 
uniform chocolate-brown colour. 

HoLOTHURiA ALBiDA. (Plate XVI. fig. 6.) 

Body elongated, tapering somewhat at either end ; tentacles darker 
(? twenty) ; suckers sparse, scattered. Body-wall thin ; oesophageal 
ring very feeble; Polian vesicle double; lungs poorly developed. 
The specimen examined had no genital tubes. 

The largest specimen was 180 millim. long, had a greatest width of 
35 miUim., and was 18 millim. wide in the region of the anus. 

The flat plates are very irregular in form ; the turriform bodies 
have a single connecting bar, and are knobbed at the narrower end, or 
where the bar is developed ; at the wider end there are also knobs, 
and these are surrounded by rather coarse spines (Plate XVI. fig. 6). 

The position of this species in the keys of Lampert cannot as 
yet be exactly determined, owing to the retracted condition of the 
tentacles ; it clearly belongs to the group of " Aspidochirote Formen 
mit Stiihlchen und glatteu Schnallen," and those in which the 
Schnallen are irregular. Like H. immobilis, it has two Polian 
vesicles ; but it difiers in colour, in the arrangement of its suckers, 
and the form of its spicules. Like H. pardalis, it has a number of 
regular plates, but it wants the characteristic marking of that species, 
and is of much larger size than any known examples. 

HoLOTHURiA CADELLi. (Plate XVI. fig. 7.) 

Body rounded, tapering posteriorly, dark above, lighter below ; 
prickly papillse, not so numerous or prominent as in U. squamifera, 
frequently but not always with a white circular base. Body-wall 
thin. 

Tentacles (retracted) ; oesophageal ring small and inconspicuous, 
the anterior region so contracted that the disposition of the Polian 
vesicles cannot be certainly made out. Cuvierian organs in the 
form of rather numerous stout csecal tubes from half to one inch in 
length. Lungs well developed. Genital tubes numerous and ex- 
tensive. 

The flattened spicules are (Plate XVI. fig. 7) of the type of those 
found in H. albiventer, but the knobs are more numerous, and 



1887.] FROM THK ANDAMAN ISLANDS. 145 

there are more than tlnee pairs of lioles ; from that species it is 
to be at once distingnislieJ by the absence of the remarkably modi- 
fied turciform spicules. On the whole, it stands nearest to //. scabra, 
but is distinguished by its speckled and less dense integument, and 
the absence of the median ventral groove. A specimen ICO millim. 
long has the greatest circumference 40; one 140 milhm. is only 10 
millim. round at the anus. 

Three smaller specinsens differ from the more matured, two by a 
larger amount of orange in their coloration, and the third by the 
greater stiffness of the skin. 

HOLOTHURIA MARMORATA. 

I have had great difficulty in coming to a definite conclusion as 
to the name to be given to the specimens now assoriated as H. inar- 
morata ; the complete absence of a circular disposition of the 
pigmentation prevents their association with H. aryus ; on the other 
hand, the comparatively small size of the oesophageal ring is against 
their affinity with H. marniorata. I believe the fact of the matter 
is that Dr. Theel is justified in his supposition that these two species 
and some others are but varieties or various stages of a variable aud 
widely distributed species which grows to a great size. 

HoLOTHURIA PAPILLATA. (Plate XVI. fig. 8.) 

Twenty tentacles. Body elongated, may be wider iu its hinder 
than in its two anterior thirds ; prominent scattered dorsal papillge, 
which are larger and more closely jiacked anteriorly than posteriorly ; 
five or more small pajjillio around the anus. 

Sucliers on central suiface only, arranged in two irregular rows, 
which unite posteriorly ; each sucker is placed on a yellowish papil- 
liform process. Colour dark slate-grey above, lighter below. 

Body-wall thin, the parts of the oesophageal ring are small, the 
ampullae long ; one large Polian vesicle. Genital tubes short and 
numerous. Apparently no Cuvierian organs. The only calcareous 
deposits are iu the form of stools (Plate XVI. fig. 8). 

Notwithstanding these numerous negative characters, the large 
size and well-developed papillae must make this a very conspicuous 
species. 

It may be 280, 320 long, and 60, 80 millim. broad. 

EXPLANATION OF PLATE XVI. 

Figs. 1-3. Moimstygia, X U. 

Fig. L A^'ith spines, and from the side. 

2. Test denuded, to show flie arrangement of the plates. 

3. Test from above, to sliovv the disposition of the ambulacra . 

4. Spicules of Haplodactijla avdamanensis. 
6. Spicule of PgcudocM-utnis ackula. 

6. Spicules of Holothuria albida. 

7. Spicules oi Holothuria cadelli. 

8. Spicules oi Holothuria papillata. 
Figs. 4-8 X 500. 

Proc. Zool. Soc— 1887, No. X. 10 



146 MR. G. A. BOULENGER ON REPTILES AND [Feb. 15, 

3. On a Collection of Reptiles and Batracliians made by 
Mr. H. Pryer in the Loo Choo Islands. By G. A. 

BoTJLENGERj F.Z.S. 

[Eeceived January 28, 1887.] 

(Plates XVII. et XVIII.) 

Our Corresponding ]\Iember Mr. H. Pryer has presented to the 
British Museum an important series of Reptiles and Batraehians, 
formed by him during a recent visit to the Loo Choo Islands. This 
being the first herpetological collection from that group of islands that 
has reached Europe, its interest in exceptionally great. The North- 
Pacific Exploring Expedition, iinderCapt. J. Rogers, U.S.N., visited 
the Loo Choo Islands in 18-54-55, and a collection of Reptiles was 
made, upon which Hallowell reported (Proc. Acad. Philad. I SCO, 
p. 490 et seq.). Unfortunately, owing to the imperfection of his 
descriptions, little more than a score of new names was gained 
by that author's contribution. Now that actual specimens are 
at hand, identification of most of Hallowell's specimens has become 
possible ; and it is my pleasant duty to furnish diagnoses of those 
species together with a list of the others collected by Mr. Pryer, 
among which are two entirely new. 

REPTILIA. 
Lacertilia. 

1. Gecko japonicus, D. & B. 

? Hemidactylvs marmoratus. Hallow. I. c. p. 491. 
? Hemidactylus inornatus. Hallow. /. c. p. 492. 

2. Ptychozoon homalocephalum, Crev. 

The unique specimen in the collection, a male, though in other 
respects agreeing with the Indo-Malayan specimens, is remarkable 
for having only eight prseanal pores. 

3. Japalxjra polygonata. Hallow. (Plate XVII. fig. 1.) 

Biploderma pohjgonatum. Hallow. /. c. p. 490. 

As was to be expected, this species is closely allied to J. swinhonis, 
Gthr. It differs in having the tibia constantly shorter than the skull, 
fewer labials as a rule, and in coloration. The number of upper and 
of lower labials is six or seven, very rarely eight. The hind limb, 
stretched forwards, reaches between the commissure of the mouth 
and the anterior border of the orbit. Adult yellowish green above, 
the interspaces between the scales black ; throat and nuchal crest of 
males bright yellow ; females with broad black cross bands on the 
hack ; tail, in both sexes, annulate with black ; young more greyish, 
with black cross bands on the back. 



p. Z S. 1887. pixvn. 





p. Smit. Qfil . et lith . 



Minterrv Bros. imp. 



1. JAPALURA POLYGONATA. 

E. TACHYDROMUS SMAHAGDINUS. 



P.Z.S. 1887. Pl.IVffi. 









Ppter Snul. del et)itX. 



Mmtem Bros . in^ . 



1. TACHYDROMUS SMARAGDINUS. 
2. LYGOSOMA PELLOPLEURUM. 3 . TROPID ONOTUS PRYER:. 




K'l^jc.. 



..., vW" 



^<^/ 

.^' 



1887.] BATRACHIANS FROM THE LOO CHOO ISLANDS. 147 

milliin. millim. 

Total length., 292 215 

Head 24 18 

Width ofhead 15 12 

Body 58 50 

Fore limb 38 33 

Hind limb 65 52 

Tibia 20 16 

Tail 210 147 

The species is represented by numerous specimens in Mr. Fryer's 
coliectiou. 

4. Tachydromus smaragdinus, sp. n. (Flate XVII. fig. 2, 
and Flate XVIII. fig. 1.) 

Head long as in T. sexUneatus, its width being contained nearly 
twice in its length. A series of granules, usually incomplete, between 
the supraoculars and the supraciliaries ; a small shield usually (19 
cases out of 20) separates the large anterior supraocular from tbe 
loreal ; temporal scales obtusely keeled, much smaller than in the 
other species, 10 to 15 on a line between the orbit and the tympanum. 
Dorsal shields strongly keeled, in eight to ten longitudinal series, 
the three outer series on each side largest and equal ; eight longi- 
tudinal series of large, strongly keeled, mucronate ventrals, and 
three or four series of smaller ones on each side. Frseanal shield 
usually longitudinally divided in the female, entire in the male. 
One inguinal pore on each side. Emerald-green above ; a pale 
yellow streak along each side, from the upper lip to the groin ; 
males usually with a broad grey or bronzy lateral band above the 
yellow streak ; lower surfaces greenish yellow. 

6. 2. 

millim. millim. 

Total length 187 207 

Head 12 13 

Width of head 6-5 7 

Body 38 42 

Fore limb 20 20 

Hind hmb 27 27 

Tail 137 152 

Numerous specimens. 

5. Lygosoma pellopleurtjm, Hallow. (Plate XVIII. fig. 2.) 

Lygosaurus pellojjleiirvs, Hallow. /. c. p. 496. 

Body elongate, limbs short ; the distance between the end of the 
snout and the fore limb is contained twice in the distance between 
axilla and groin. Snout very short, obtusely acuminate. Lower 
eyelid scaly. Nostril pierced in a single nasal ; no supranasal ; 
frontonasal broader than long, forming a broad suture with the 

10* 



148 MR. G. A. BOXJLENGER ON REPflLES AND [Feb. 15, 

rostral and a still broader one with the frontal ; prtefrontals very 
small and widely separated ; frontal short, in contact with the first 
supraocular only, followed by a long and narrow single frontoparietal 
resembling a second frontal ^ angular posteriorly and in contact 
with the interparietal; latter in the middle between four small 
parietals, the posterior pair of which are in contact behind ; four 
supraoculars, first in contact with the frontal and the frontoparietal, 
second and third in contact with the frontoparietal ; seven supra- 
ciliaries ; fourth upper labial below the centre of the eye. Ear- 
opening small, horizontally oval, without projecting lobules. 24 
scales round the middle of the body, subequal; dorsals rather strongly, 
laterals feebly tri- or qninquecarinate. Prseanal scales not enlarged. 
Limbs widely separated when adpressed ; the length of the hind 
limb equals the distance between the fore limb and the nostril. 
Digits short, cylindrical : subdigital lamellse smooth, 10 under the 
fourth toe. Tail a little longer than head and body. Reddish 
brown above, dark brown on the sides ; a black lateral band, com- 
mencing from the eye; sides with fine whitish shafts; belly dirty 
white, spotted with dark brown ; lowei- surfaces of tail plumbeous 
grey, spotted with black. 

millira. 

Total length 12.5 

Head 9 

Width of head &-5 

Body 48 

Fore limb 10 

Hind limb :.... 16 

Tail 68 

A single specimen is in the collection. 

O P H I D I A. 

6. Ablabes semicarinatus. Hallow. 

Eurypliolis semicarinatus, Hallow. /. c. p. 493. 

Scales in fifteen rows, without apical groove, with a very feeble 
keel along their anterior half. Rostral rather broader than deep ; 
nostril between two nasals ; prcefrontals about twice as long as 
internasals ; frontal much longer than broad, as long as or slightly 
longer than internasals and praefrontals together ; loreal at least 
twice as long as deep ; one prte- and two postoculars ; eight upper 
labials, fourth and fifth entering the eye, seventh largest ; temporals 
1+2 ; five lower labials in contact with gulars. Ventrals 187-192; 
anal divided ; subcaudals 72-82. Olive above, the scales lighter in 
the centre ; the lower scale on each side yellowish, margined with 
olive or black ; in some specimens four rather indistinct dark brown 

^ The very singular shape of this shield misled Hallowell as to the homo- 
logies of the head-shields of this Lizard. Therefore his interuasal = fronto- 
nasal ; frontonasal = I'rontal ; frontal = frontoparietal ; interparieto-frouto- 
parietal — interparietal ; frontoparietals = anterior parietals. 



1887.] BATRACHIANS FROM THE LOO CHOO ISLANDS. 149 

longitudinal hands on the hinder half of the hody ; tipper lip, ventrals, 
and subcaudals uniform yellowish. 

Several specimens, the largest measuring T] centim. 

7. Tropidonotus pryeri, sp. n. (Plate XVIII. fig. 3.) 
Dentition syncranterian. Form slender, as in T. saurita. Tail 

one third of the total length. Head and eye moderate. Scales 
in 19 rows, all strongly keeled. Ventrals 17(5-180 ; subcaudals 
121-128. Internasals truncated in front, more than half as large 
as the prsefrontals ; one prae- and three postoculars ; loreal a little 
broader than deep ; normally eight upper labials, fourth and fifth 
entering the orbit ; usually two temporals in the first row. Anterior 
part of body with large alternating black elongate spots, separated 
by a narrow zone of pale olive on the vertebral line, by broader 
yellowish vertical bars on the sides. Head dark olive, above spotted 
or marbled with black ; lip yellowish, the sutures between the 
anterior upper labials black ; an oblique black streak across the 
(normally) seventh upper labial ; a yellow spot, surrounded with 
black at the extremity of the jaws. Hinder part of body with a 
series of black dorsal spots and a lateral series of light spots. Lower 
parts yellowish, the hinder part of tlie body with a series of black 
streaks along each side of the ventrals, becoming confluent into a 
lateral black line on the subcaudals. 

Four specimens, the largest measuring 93 centim. The stomach 
of one contained a Rana gracilis. 

8. Platurus fasciatus, Daud. 

9. BoTHROPS FLAvoviRiDis, Hallow. /. c. p. 492. 

Upper and lower head-scales smooth ; upper very small, seven or 
eight in a transverse series between the supraoculars, which are 
large ; nas:d divided ; eight upper labials, third and iburth largest, 
second borderirig the loreal pit. Scales small, 35 across the middle 
of the body ; laterals smooth, the others feebly keeled. Ventrals 
230 ; anal entire ; subcaudals, 75 pairs. Yellowish green above, 
with symnietrical blackish markings ; lower parts pale yellow, with 
pale olive spots confluent into two bauds on the anterior part of the 
belly. 

A single specimen, 525 millim. long. 

BATRACHIA. 

Ec AU DATA. 

1. Rana GRACILIS, Wiegm. 

2. Rhacophorus viridis. Hallow. 
Polypedates viridis, Hallow. I. c. p. 500. 
? Hyla cyanea. Hallow. I. c. p. 494. 

Closely allied to i?. Sihlegeli, Gthr., from which it differs in the 
longer hind limb, the tibia measuring half the length of head and 
body ; when the limb is stretched forwards, the tibio-tarsal articulation 



150 MR. O. THOMAS ON SMALL [Feb. 15, 

reaches between the eye and the nostril. The Inner metatarsal 
tubercle is smaller than in li. schlegeli, and quite flat. 

A single specimen, measuring 53 millim. from snout to vent. 

3. MiCROHYLA FISSIPES, Blgr. 

C AU DATA. 

4. MOLGE PYRRHOGASTRA, Boic, Var. ENSICAUDA, HalloW. 

Triton subcristatus (ensicauda), Hallow. I. c. p. 494. 

These specimens (ten in number) appear sufficiently different from 
the Japanese typical form to warrant a varietal distinction. The 
head is proportionally somewhat broader, the digits shorter, and the 
tail, in the female, longer. The lower parts are yellow (not red), 
immaculate or more or less spotted with black ; digits yellow 
inferiorly (black in the typical form). Upper parts black, in some 
specimens with scattered small yellowish or pale green spots ; 
sometimes a yellowish vertebral streak. 

c?. $. 

millim. millim. 

Total length 115 158 

From snout to cloaca 53 70 

Head 13 15 

Width of head 12 14-5 

Fore limb 20 21 

Hind limb 20 23 

Tail 62 88 

EXPLA]!^ATION OP THE PLATES. 
Plate XVII. 

Fig. 1. Japalura folygonata, p. 146. 

2. Tachydronms smaragdinus, ji. 147. 

Plate XVIII. 

Fig. 1. Tachydromus smaragdinus. Upper yiew of head, X 2. 

2. L)/gosoma peUojdi'urum ; and upper view of head, X 2, p. 147. 

3. Trojjidonotus prycri, p. 149. 



3. Oil the small Mammalia collected iu Demerara by 
Mr. W. L. Sclater. By Oldfield Thomas. 

[Eeceived January 29, 1887-] 

(Plate XIX.) 

The Mammalia obtained by Mr. W. L. Sclater during his recent 
visit to British Guiana consist of 13 specimens belonging to 8 
species, of which one is new. All of them have had their exact 
localities and dates recorded, and are therefore of interest even when 
belonging to common species. The discovery of a new species of the 



P.X S. 18B7. PL XIX. 




:1 l<l.h 



iiliai'l. 



aiuiai'L, imp 



m>p 



HF.SPl'ROMYb IRHIPIDOMYS^ Sl'I.ATF.RI 



1887.] MAMMALS FROM DEMERARA. 151 

peculiar Dormouse-like subgenus Rhipidomys is a very interesting fact, 
and shows how much these small mammals have been neglected by the 
very numerous collectors who have worked in the different parts of 
Guiana. The following is a list of the species, with short notes by 
Mr. Sclater upon their habits, &c. 

1. Vesperugo (Vesperus) hilarii, Geof. 

a. Maccasseema, Pomeroou R., 15/12/86. 
" Caught in the store-room in the daytime ; apparently it VFas 
crawling about the place, and made no attempt to fly away." — 
W. L. S. 

2. Furia horrens, F. Cuv, 

a. Maccasseema, 11/86. 
"This was the common House-bat of Maccasseema and flew 
about the premises at dusk. The individual obtained was killed by 
Mr. imThurn."— ir.i. /S'. 

3. Rhynchonycteris naso, Wied. 

a~d. Marakka, 20 miles up the Pomeroon, 15/12/86. e. Near 
Maccasseema, 5/12/86. 
•' These Bats cling most of the day to the stump of a tree over- 
hanging the water, and when disturbed take short flights and again 
return to the same place." — W. L. S. 

See also Dobson, Cat. Chiropt. B. M. p. 368 (1878), and ira 
Thurn, 'Among the Indians of Guiana,' p. 115 (188J). 

4. Saccopteryx leptura, Schr. 

a. Calicoon, close to the junction of the Essequibo and Maza- 
runi Rivers, 14/11/86. 

"This Bat I knocked down about dusk, say 5.30 p.m., with a 
long stick, just outside the house. Several others were flying about, 
but I was unable to secure any more." — 7F. L. S. 

In this specimen the wing-pouches are unusually large and dis- 
tended, and from each of them there projects a prominent white 
frill of skin, which can apparently be exserted or withdrawn at 
pleasure. From the marked development of these organs, un- 
doubtedly sexual in their nature, it may be inferred that the late 
autumn is the pairing-time of this species, at least in Guiana. 

5. Glossophaga soricina. Pall. 

a, b. Pen Hope, 13/10/86. On the coast 20 miles east of 
Georgetown. 
" This Bat was found in the house in considerable numbers." — 
W. L. S. 

6. HoLOCHiLus (Nectomys) sauAMiPES, Brandt. 

G.Young. Pen Hope, 12/10/86. 
"Caught in the cane-fields, and brought to me by a coolie." — 
W. L. S. 



152 ON SMALL MAMMALS FROM DEMERARA. [Feb. 15, 

7. Hesperomys (Rhipidomys) sclateri, sp. n. (Plate XIX.) 

a. 5 . Maccasseema. 1 1/86. 

"I am sorry to say I caunot remember anything about the habits 
of this Rat ; it was caught aud brought to me by one of Mr. im 
Thurn's Indians, and of course did not live in tlie house." 

Fur short, close, very soft and velvety. General colour uniform dark 
ashy grey, the tijis of the hairs below white or pale rufous, line of 
demarcation not strongly marked ; bases of all the hairs slate- 
coloured. Hairs on both fore aud hind feet, including the fingers 
and toes, all dark brown or black. Ears, when laid forward, reaching 
just to the centre of the eye ; no projection on their anterior border ; 
tlieir backs hairy, black. Tail long, uniformly black, thickly hairy, 
the hairs about 3 or 4 mm. long throughout, except just at the base, 
where they are shorter, and at the extreme tip, where they are 10 or 
12 mm. long; the rings of scales well-marked, 15 or 16 to the 
centimetre. INIammse 6, one axillary and two inguinal pairs. Inter- 
dental palate-ridges 6. Foot-pads broad, smooth, rounded ; soles 
naked, quite smooth. 

Skull exceedingly similar to that of H. leucodactylus, Tscb. 
(figured P. Z. S. 1884, pi. xliv. fig. 8), but rather longer and nar- 
rower, especially in the cranial portion, with the supraorbital edges 
more strongly developed, and with the incisors rather longer and 
heavier. 

Dimensions of the type, an adult female in spirit : — Head and 
body 133 mm., tail 1/2, hind foot 33, forearm and hand 39, ear, 
above crown, 16, head 43, muzzle to eye 18'5. 

Skull. Basal length 31-5, greatest breadth 19; nasals, length 
12-8; length of molar series 6-4; back of incisors to front of 
m^ 102; palatine foramen, length 8-0; interorbital constriction 
6-3. 

This species, with which I am glad to connect the name of its dis- 
coverer and donor, is very closely allied, in all its essential characters, 
to H. (^Rhipidomys) leucodactylus, Tscb.-, but that s])ecies has its fur 
very much the nature, colour, and texture of that of the Common 
Rat ; while in //. sclateri tlie fur is wholly difTerent to this, being in 
tact more like that of certain of the smaller Opossums in its soft and 
velvety character. In //. sclateri the colour is also darker and more 
uniform than in H. leucodactylus, the tail is more uniformly busliy, 
and the feet, both fore and hind, differ by having black-haired 
instead of pure white toes. 

The present is the first recorded occurrence of any member of the 
interesting Dormouse-like subgenus Rhipidomys in the region north 
of the Amazons and east of Colombia, and gives therefore a very 
important addition to the known range of the subgenus. Other 
species have beeu recorded from Central America, Ecuador, Peru, 
Bahia, and jNIinas Geraes, the nearest ally of //. sclateri being, as 
already noted, the Peruvian H. leucodactylus, Tscb. 

2 Fauu. Peruana, p. 183, pi. xiii. fig. 2 (1844). 



1887.] MR. BOULENGER ON A NEW GECKOID LIZARD. 153 

8. DiDELPHYS MURINA, LlUn. 

a. Pen Hope. 12/10/8G. 
"Brought to me by a coolie at Plantation Hope, which is on the 
so-called east coast, i. e. about 20 miles along the coast east of 
Geori>;etown ; it was caught, as I understand, in the cane-fields." — 
JF. L. S. 



4. On a new Geckoid Lizard from British Guiana. 
By G. A. BouLENGEK, F.Z.S. 

[Eeceived January 24, 1887.] 

A small collection of Pteptiles and Batrachians was formed by Mr. 
W. L. Sclater during his recent visit to Maccasseema (on the Pome- 
roon River) in British Guiana, and presented to the Natural History 
Museum. Small as the collection is, and from a comparatively well- 
explored district, it nevertheless contains a novelty, the small Lizard 
described below. The other species of which specimens were 
obtained are the following : — • 

Lizards : Thecadaetijlus rapicauda, Houtt. ; Anolis punctatus, 
Daud.; OphryoessasvpercUiosa,'L.; Uraniscodun umbra,!,.; Cophias 
Jiavescens, Bonn. ; Amphisbcena fuliginosa, L. 





Gonatodes annularis. 



154 MR. r. E. BEDDARD ON A [Feb. 15, 

Snakes : Typhlops reticulalus, L. ; Geophis lineatus, D. &, B. ; 
Elaps lemniscatus, L, 

Batrachians : Leptodactylus pentadactylus, Laiir. ; Sufo marinus, 
L. ; Bufo typhonius, L. ; Coecilia gracilis, L. 

GONATODES ANNULARIS, Sp. n. (Woodcut, p. 153.) 

Closely allied to G. albogularis, D. & B. Snout longer than the 
diameter of the orbit, obtuse, the granules on its upper surface 
small, not larger than the dorsals. Supraciliary edge with a small 
projecting spine, as in most species oi Sphcerodactylus. Seven upper 
and four or five lower labials ; a pair of small chin-shields behind 
the mental. No transversely enlarged subcaudals. Grey-brown 
above, with a series of large black spots along each side of the 
vertebral zone ; head and limbs with black spots or marbhngs ; 
tail with black annuli, alternating with white spots inferiorly ; lower 
surfaces pale brown, throat with oblique dark-brown lines converging 
backwards. 

millim. 

Total length 86 

Head 11 

Width of head 7 

Body 29 

Fore Hmb 15 

Hind limb 19 

Tail 4G 

Two female specimens. 



5. On the Structure of a new Genus of Lumbricidse 
{Thamnoclrilus guUelmi^). By Frank E. Beddard, 
M.A., F.R.S.E,, Prosector to the Society, Lecturer on 
Biology at Guy's Hospital. 

[Eeceived February 15, 1887.] 

I owe the specimens of the worm described in the present paper to 
the kindness of Mr. W. L. Sclater, F.Z.S., who collected them for 
me during his recent visit to British Guiana. 

They are all rather large worms (see woodcut, fig. 1), measuring 
up to 6 inches in length and g inch in breadth, and belong to a new 
genus and species of Lumbricidse, which I propose to call Thamno- 

DRILUS GULIELMI. 

External Characters. — The colour is purplish on tlie dorsal 
and reddish yellow on the ventral side : the clitellum is distin- 
guished from the rest of the body by its paler tint. 

The mouth is situated precisely at the anterior extremity of the 

1 Named after Mr. William Lutley Sclater. 



1887.] 



NEW GENUS OF LUMBRICID^. 



155 



Fig. 1. 






llM 






•Vliil 



/UtPltil 







Thamnodnlusgulielmi, from the ventral surface; natural 



size. 



156 MR. F. E. BEDDARD ON A [Feb. 15, 

body, and there is therefore no prostomium ; in this character 
Thamnodrilus agrees with UrochtBta^ and Biachafa'. It is a little 
difficult to distinguish the anterior segments of the body ; the 
buccal segment is divided externally by a very well-marked furrow, 
which appears to mark the line of division between two really 
distinct somites. A consideration of the number of the nephridia 
(see p. 160) leads me to infer that the divisional furrow does not 
imply a division into somites ; furthermore (see tigs. 1 and 2) each of 
the two anterior rings is divided on each side by a longitudinal 
furrow, which corresponds in position with the dorsal pair of setse 
in the following segments ; the presence of this furrow is perhaps 
an additional argument in favour of regarding them as parts of the 
same somite. 

From the first segment to about the 8th, the breadth of the 
segments gradually increases ; the longitudinal diameter of the 
segments also gradually increases up to the 8th or 9th, after which 
they become distinctly narrower. 

The segments of the ditellum are perfectly distinct, and are eleven 
in number. The clitellum commences with the 15th segment and 
terminates upon the 25th ; very generally a trace of glandular 
modification is to be found upon the 14th and 26th segments. 

The clitelhnn of Thamnodrilus is therefore nearly coextensive with 
that of Urohenus, Urochmta, Titanus, and Anteus. 

As in the former genus, the clitellum is not developed \ipon the 
ventral side of the body, but the extent of the area left free from 
glandular substance varies ; in the anterior part of the clitellum, 
as far back as segment 19, the ventral pair of setae as nearly as 
possible mark the boundary between the glandular and non-glandular 
portion of the integument ; from the 2Gth segment to the end of 
the clitellum there is a space left between the glandular part of the 
integument and the setae. This is illustrated in the accompanying 
drawing (woodcut, fig. 2). 

The setce are disposed in pairs ; the distance separating the 
ventral pairs is 2"2 mm. in the clitellar region, the distance between 
the ventral and dorsal pair is 5"5 mm., VAhile the dorsal pairs are 
separated by an interval of 12"5 mm. The setae are not remarkable 
in shape except upon the clitellum ; here they are modified and 
acquire the form illustrated in the drawing (woodcut, fig. .3) ; these 
peculiarly modified setae are apparently found throughout the 
clitellum (I have also found them in segments immediately anterior), 
where they replace both the dorsal and ventral pairs. The accom- 
panying figure renders any minute description of these setae unneces- 
sary, and will serve to show how exactly they resemble the clitellar 
seta; of Vrochceta ; in this genus Perrier has recorded ^ the modi- 
fication of the ventral pair of setae on the 20th segment ; and Ilorst 
has stated ^ that in another species the ventral setae of four of the 

1 Perrier, Arch, de Zool. Exper. t. iii. 

^ Benham, Quart. Jouni. Micr. Sci. 1886. 

2 Arch. d. Zool. Exper. t. iii. (1S74) p. 31)9, pi. xvii. fig. 52. 
* Midden Sumatra, Verme.s, p. 8. 



1887.] 



NEW GENUS OF LUMBRICID^. 



15; 



c itel ar segments are thus modified. This latfer statement I am 
able to confirm from the study of an Austrahan species of the genus 
possibly identica with Ilorst's species. But the modification of the 
setae m the chtellar region of Thamnodrilas is more complete than 




Fig. 3. 




Pig. 2. ThamnodrilmguhelTm. Anterior segments seen lateraUy ; apertures of 
the nephridia in front of the dorsal pair of setce ; segments of clitellum 
PV q c .} ^'^'■^^''a^y- Magnified twice natural size. 
1 ig. 6. beta from one of the segments of the cliteUum; highly magnified. 

in Vrochata, inasmuch as it has affected the dorsal as well as the 
ventral pairs. The clitellar setae are larger than the ordinary setae, 
as well as differmg in their ornamentation ; this difference was very 
noticeable m an immature example, where the difference of age alone 



158 



MR. F. E. BEDDARD ON A 



[Feb. 15, 



distinguislied the clitellar from the ordinary setse ; moreover I have 
frequently found even in mature examples that the clitellar setae of 
both dorsal and ventral pairs only showed slight traces of ornamen- 
tation at their free extremity ; such setfe alternated in the most 
irregular fashion with seta3 like that displayed in the drawing (fig. 3) ; 
they are, however, of equal size. 

Dorsal i^ores appear to be entirely absent. 

The only ajjertures besides the mouth and anus recognizable on 
the exterior of the body are those of the nephridia, which are placed 
in front of the dorsal pair of settc close to the anterior margin of 
the segment (fig. 2) ; these apertures were especially plain upon the 
clitellum. 

Vascular System.— In none of my specimens was the vascular 
system very well preserved. The dorsal vessel (fig. 4, d) runs close 
to the surface of the gizzard, and near to the hinder end of that 
organ gives off two pairs of slender trunks (A) to the ventral vessel (v). 
After this the dorsal vessel runs some way above the surface of the 
oesophagus, and is not directly connected with the ventral vessel ; 
segments 10, 11, and 12 contain each a pair of lateral "hearts" {h), 




Chief Trunks of Vascular System. 

e?, the dorsal vesse] ; r, Tentral vessel ; .«, supvaiiitestinal ; ^, anterior " hearts " 
connecting dorsal and ventral vessel; h', posterior hearts connecting 
supraintestinal and ventral vessel. 

of which the two anterior pairs are considerably stouter than the pos- 
terior pair ; these are given off from the supraintestinal trunk (s), 
which is of some thickness in these segments. I could not detect 
anv additional communication with the dorsal vessel, and am inclined 
to think that there is none. 

Alimentary CanaJ. — The mouth-apertvre, as already stated, is 
terminal, and in all my specimens was excessively minute ; the buccal 
cavity is thick-walled and very narrow, with an almost imperceptible 
lumen. The pharynx is large, and is followed by a wide thin-walled 
a'sophagus. The oesophagus is remarkable for the fact that it does 
not pass straight to the gizzard, but is bent upon itself, forming a 
loop : it might be readily imagined that this condition is simply due 
to the contraction produced by the preservative reagent ; but I 
found the same condition of the oesophagus in all the specimens that 
I dissected, and in all of these the gizzard was apparently retracted 



1887.] NEW GENUS OF LUMBRXClDiE. I59 

rather than protruded, and the segments of the body in most instances 
were perfectly normal and not unduly contracted. The gizzard is hke 
that of other Earthworms. 

(Esophageal Glands. — Behind the gizzard and in front of the 
intestines the oesophagus is furnished with certain glands, which 
are evidently the homologues of similarly placed glands in other 
Earthworms. These glands are kidney-sliaped and lie on the dorsal 
or lateral aspect of the CEsophagus, with the concave side turned 
towards the oesophagus ; at the middle of its concavity the gland 
is connected with the oesophagus by a short duct, the general 
appearance of these glands is strikingly similar to the " kidney- 
shaped glands" which I have described"' in the intestinal region of 
Megascolex caruleus ; a])parently they also resemble in outward 
appearance the oesophageal glands of Notoscolex camdenensis'-, 
though frequently the CESophageal glands of Earthworms hare not 
this peculiar shape. 

The oesophageal glands of Thamnodrilus are furnished with a 
very abundant blood-supply. This blood-supply is derived from 
the supraintestinal trunk, and not from the dorsal vessel ; in the 
case of the two posterior pairs, at any rate, of these glands, a 
branch is given off on either side from the supraintestinal vessel ; 
this at once divides into two trunks ; the inner branch goes to 
the septum and ramifies upon its surface; the outer branch 
conveys blood to the oesophageal gland, which it reaches by 
passing along the pedicle by which that gland is attached to the 
oesophageal walls ; the vessel then breaks up into a network of 
capillaries on both the anterior and posterior surfaces of the o^land. 
It is important to notice that in these segments both the dorsai 
region of the mesentery and the oesophageal gland are supplied with 
blood from the supraintestinal trunk ; the dorsal vessel o^ives off 
no branches in these segments. In Urochceta the calciferous glands 
have, according to Perrier, a similar blood-supply. 

There were altogether six pairs of these glands situated in seg- 
ments 9-14; the last two pairs, i. e. those situated in segments 
13 and 14, were situated nearer to the dorsal surface of the in- 
testine than those which preceed them. The number of those 
glands (six pairs) is unusual, three being the almost constant number 
of pairs in other Earthworms; in certain species of Perichceta, 
however, there appear to be as many as six pairs of cesophaoeal 
glands. ® 

Body-cavihj.—lhe dissection of this part of the body was rendered 
very difficult by the toughness of the septa in this region and by their 
firm connection with one another by numerous tendinous threads; 
these septa, however, in Thavmodrilus, are not specially thickened^ 
as they are in many other Earthworms, but are thin and "transparent' 
as in the posterior region of the body. * 

The body communicates with the exterior only by the apertures 
of the uephridia ; there are no dorsal pores present. 
' Tran.s. Eoy. Soc. Edinb. vol. xxxii. 
^ Proc. Linu. Soc. N. S. W. 1886, pi. viii. fig. 1, i.g. 



160 



MR. F. E. BEDDARD ON A 



[Feb. 15, 



Nephridia. — The nepliridia appear to be present in all the 
segments of the body with the exception of the first ; they are 
differentiated into three series. 

(1) The first series consists of only one pair of nephridia ; these 
differ from those which follow in their structure and in tlieir posi- 
tion. They lie beneath the oesophagus and are completely hidden 
by it ; each gland consists of a flattened mass of glandular tubules, 
produced by the coil, which has the ordinary structure characteristic 



Fig. 5. 




One of the Anterior Nej)bridia. 

0, external aperture; f, funnel opening on to the other side of the 
intersegmental septum s ; d, glandular Tcsicle. 



of nephridial tubules, except that the coils are more numerous. The 
tubule opens into a stout-walled muscirlar duct, distinguishable by 
its yellowish colour, which passes anteriorly in a somewhat sinuous 
course and opens on to the second segment of the body. These 
glands evidently correspond to the " glandes a mucosite " described 
by Perrier in Urochceta^, and by myself in AcanthodrilusmuJtiporus-, 
The funnel was very conspicuous in tiansverse sections. This pair 
of nephridia differs less from the succeeding pairs than in many 
other genera. The specialization of this first pair of nephridia. 



1 Arch, de Zool. Exp. t. iii. 



s P. Z. S. 1885. 



1887.] 



NEW GENUS OF LUMBRIC1D.E. 



161 



which is so well marked ia JJrochceta and ia Acanthodrilus multi- 
porus, is, as it were, just commencing in Tkamnodrilus. 

(2) The next fourteen pairs of nephridia (see fig. 5) open on to the 
exterior of the body in a line with the dorsal pair of setfe ; the most 
anterior nephridia are rather smaller than the posterior pairs in corre- 
sponding with the increasing width of the segments. The glandular 
part of the nephridium is very slightly developed in conijiarison with 
the extremely elongated muscular sac which communicates with the 
exterior ; at the junction of the two regions of the nephridium, the 

Fig. 6. 




One of the Posterior Nephridia. Lettering as in fig. 5. 

muscular duct becomes dilated and bent slightly upon itself; its 
walls become glandular ; this portion of the gland is distinguishable 
by its opaque white appearance. 

(3) From the 17th segment to the end of the body the nephridia 
are of a somewhat different character to those which precede 
them ; the muscular sac of the gland is well developed and opens 
on to the exterior at the same point as in the anterior segments ; 
it differs in being furnished with a diverticulum which is nearly as 
long as itself; the glandular region of the nephridium is more 

Proc. ZooL. Soc— 1887, No. XI. H 



162 ON A NEW GENUS OF LUMBIIICID.E. [Feb. 15, 

complicated in the first few segments after the 17th than in the 
posterior segments of the body ; the proximal end of the muscular 
duct passes into a somewhat dilated pear-shaped glandular vessel, 
into which opens the nephridial tubule ; in the posterior nephridium 
(see fig. 6) the nephridial tubule is very short and hent upon 
itself four times, the four tubes running parallel veith each other ; 
at a point about opposite to the glandular vesicle the tubule perforates 
the mesentery and reaches the interior of the segment lying in front 
of that which contains the distal part of the organ ; here it ends 
almost immediately in the nephridial funnel, which is very large and 
conspicuous ; instead of being a simple funnel-shaped expansion, as 
in the majority of Earthworms and in the anterior nephridia of this 
species, this region of the nephridium forms an elongated folded 
membrane apparently closely agreeing with the nephridial funnel 
of Anteus^ ; this membrane is composed of the ordinary columnar 
ciliated cells. 

Reprodiietive Organs. — There are two pairs of vesiciiIcB seniinales, 
situated in segments 11 and 12; each of these bodies is somewhat 
kidney-shaped and flattened laterally ; the membrane covering the 
vesiculse is continued over the funnels of the vasa deferentia which 
open into the same segments. The true testes were conspicuous in 
a young specimen which I investigated by means of transverse 
sections. 

I traced back the vasa deferentia, as an excessively fine tube, as 
far back as the 18th segment, where it appears to open close to the 
ventral median line. I could not, however, detect the actual orifices 
of the vasa deferentia. Prostate glands appear to be entirely absent, 
as in many (e.g. Urocho'ta, Microcliata) of Perrier's Intraclitellians ; 
this grouf), however, cannot be distinguished by the absence of 
prostate glands, which are present in Eudrilus, JMec/ascolex, and 
Typhcpus. The ovaries are situated in the usual position in segment 
13 ; they are small digitate glands. 

The oviducts open by expanded funnels into the 13th segment 
which are placed close to the nerve-cord ; their ducts perforate the 
mesentery and open on to the exterior in the 14th segment. I did 
not, however, observe the actual orifice, which must be well within 
the ventral pairs of setae, if not actually unpaired and median. 

Sperm athecce. — I opened one or two mature individuals and failed 
to find these organ's ; they are, however, usually })resent to the 
number of a single pair in tlie seventh segment. Each sperma- 
theca is a simple, splierical, or pear-shaped pouch without any 
diverticulum ; it opens exactly in front of the nephridium of the 
same segment. 

It appeared to me at first that this Earthworm might possibly 
belong to the genus Aniens, E. P. ^ 

Perrier's dcscrijition of the genus is not very complete, as it 
necessarily depended upon a unique example which could be only 
partially dissected. 

' Perrier, Nouv. Arch. d. Mus. t. viii. pi. i. fig. 14. 

^ " Eecherches," &c. Nouv. Arch. d. Mus. t. viii. p. 49. 



188".] ON A NEW PARASITIC DIPTEROUS INSECT. 163 

Anteus agrees with Thamnodrilus in the absence of dorsal pores, 
in the arrangement of the setae, the position of the nephridiopores, 
the characters of their internal funnel, and in the presence of a single 
pair of spermathecaj in the 7th segment. Both genera have two 
pairs of vesiculpG seminales in segments 11 and 12. The main 
external points of difference appear to be in the clitellum, which is 
much more extensive in Anteus than in Thamnodrilus, and in the 
modification of the clitellar setse in the latter genns. In Thamno- 
drilus the anterior mesenteries are not specially thickened as 
they are in Anteus and there is no modification of the nephridia in 
the genital segments. Whatever may be the way in which the 
genital products are carried off in Anteus, the genital ducts of 
Thamnodrilus are perfectly normal. Anteus, like Thamnodrilus, is 
a native of the northern part of the South-American continent. 



6. Note on a new Parasitic Dipterous Insect of the Family 
Hippoboscidse. By Charles O. Waterhouse. 

[Eeceived January 31, 1887.] 

The insect here described was found by Dr. R. W. Shufeldt^ at 
Fort Wingate, New Mexico, on a species of Swift (Cypselus melano- 
leucus), and transmitted to Mr. Sclater for examination. It is closely 
allied to Anapera pallida, a European Dipterous parasite found on 




Cypselus apus. It is, however, much larger, and is at once distin- 
guished by tlie almost total absence of wings — a character which 

' See Dr. Shufeldt's paper on this Swift, 'Ibis,' 1887, p. 151. 

11* 



164 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

might, by some, be considered of generic importance. Having only 
two examples, which appear to be females, I prefer for the present 
to place the species in the genus Anapera, and to name it 

Anapera fimbriata. 

Smoky yellow, with the abdomen brown ; the epistome pale 
yellow. The general form and structure are those of A. jxillida, but 
it is considerably larger. The antennae are beset with long black 
hairs. There is a thick fringe of long black erect hairs or setae in 
front of the eye, continued posteriorly along the orbits of the eyes 
on each side of the middle opaque disk. This fringe exists, but in 
a much less degree, in A. pallida. The triangle on the vertex is 
longer than broad, and not transverse as in A. pallida. There is a 
series of black setae along the posterior margin of the head. The 
thorax is of the same form and with the same black setas as in A. 
pallida, but they are stronger and more conspicuous. The rudimen- 
tary wings are pale smoky yellowish, about as long as broad, with 
numerous black setae on the costal area. The abdomen is somewhat 
round, clothed with black hair, which is very short on the disk, long at 
the sides and apex ; the base has a transverse arcuate fold ; the disk 
is deeply impressed, but, although this is nearly the same in both 
examples, it is possibly (he result of contraction. The legs are as in 
A. pallida, beset with black hairs. 

Length 5 lines. 



7. On the Terrestrial Mollusks of the Viti Islands. — Part I. 
By Andrew Garrett, of Hualiine, Society Islands. 
(Commnnicated by Mr. John H. Ponsonby, F.Z.S.) 

[Received December 8, 1886.] 

The Viti Archipelago, which comprises nearly 200 islands and 
islets, is embraced in an area between 178° 20' W. and 176° 55' E. 
long., and between 15° 4/' and 19° 13' S. lat. The islands are dis- 
posed in three groups — the eastern, intermediate, and western. The 
former, which is only partially explored, comprises many small islands, 
mostly of coralline formation, which have been more or less upheaved 
through volcanic agencies. All the land-shells, so far as known, com- 
prise the same genera of small shells as obtained in the Tonga and 
Samoa Islands. The middle portion, which includes all the large 
islands, though imperfectly explored, have so far yielded many large 
and interesting species. Besides the same genera which occur in the 
eastern group, we find the genera Placostylus, Nanina, Diplommatina, 
Pupina, and Lagocheilus. All these genera, which are represented 
by peculiar species, connect the land-shell fauna with Australasia 
and the East Indies. The latest and most interesting discoveiy is 
the occurrence of the Asiatic genus Lagocheilus, which was found 
by Mr. Liardet in Gomea Island. The western or Assawa group. 



1H87.] MOLLUSKS OF THE VITl ISLANDS. 165 

which is unexplored, will undoubtedly produce many species not 
found in other parts of the archipelago. 

The genus Succinea, so widely diffused throughout the Pacific 
islands, has not been discovered in these islands. 

The earliest known Vitian endemic land-shells are Bulimus mallea- 
tus, B. fulffuratiis, and Helix noiileti, all described in the 'Revue 
Zoologique ' in 1842, the two former by Dr. Jay, and the latter by 
Le Giiillou. In 1845, Philippi published Helix pfeifferi, and Mr. 
Hinds described Pythia pollex. In 1846 and 1847, Dr. Gould 
added several new species, all discovered by the TJ. S. Exploring 
Expedition. In 1855 Dr. Pfeiffer described Helix ludersi; and six 
years later Dr. Dohrn added the beautiful Bulimus seemanni to the 
list of endemic species. In 1865, Prof. Mousson published, in the 
'Journal de Conchyliologie,' a complete list of the Viti land and 
freshwater shells, based on the collections made by Dr. GrJifife and 
added several new species to the 16 peculiar to the group. In 1870 
Mousson's second paper appeared in the same Journal, and he de- 
scribed 26 new species, all collected by Dr. Graffe. In the meantime 
several new species were published by H. Adams, Crosse, Angas, 
and Semper. 

In the ' American Journal of Conchojogy ' for 1871, and in the 
' Proceedings of the x\cademy of Natural Sciences of Philadelphia ' 
for 1873, the writer published descriptions and figures of 20 new 
species, all personally collected. The latest discovery, so far as I 
can ascertain, is six new minute species collected by Mr. Liardet, 
which were published and figured with their animals in the 'Proceed- 
ings' of the Zoological Society for 1876. 

Out of 146 species now recorded 85 are peculiar to the group. The 
146 species are embraced in 32 genera, 11 of which are operculated. 

Genus Helicarion, Ferussae. 

1. Helicarion vitrinina. 

Naninal vitrinina, Liardet, Proc. Zool. Soc. 1876, p. 100, pi. 5. 
figs. 2, 2a. 

" Shell yellow, thin, translucent, perforated, discoidal ; whorls A\, 
last subangulated, the others slightly convex ; beneath shining and 
well rounded ; aperture slightly oblique, lunate ; suture marginate ; 
peristome thin ; columella slightly expanded over the perforation. 
Animal black, with mantle covering two thirds of the shell, which it 
cannot enter at first." 

" Found in moist situations under logs, in this respect resembling 
Vitrina." (^Liardet.) 

This species, which is unknown to me, was found at Taviuni Island. 

2. Helicarion ramsayi. 

Nanina ? ramsayi, Liardet, Proc. Zool. Soc. 1876, p. 100, pi. 5. 
fig. 3. 

" Shell similar to N. ? vitrinina. When the animal is out I can 
detect no difference. 



166 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

" Animal red ; a protuberance on the back rests against the shell 
anteriorly. It progresses by raising its head, extending the body, 
and placing the posterior part of the foot down in the form of an 
arch, lands its body gradually from the head ; and this arch thus 
appears to recede until the caudal extremity is reached. Like 
N. ? vitrinina, it cannot at first recede into its shell ; and like 
Vitrina strancjei of Au'^tralia, it leaves mucus in its track of a 
brick-red colour. Ilab. Taviuni." {Liardet.) 

Genus Nanina, Gray. 

1. Nanina nouleti. 

Helix nouleti, Le Guillou, Rev. Zool. 1842, p. 137; Pfeiffer, 
Mon. Hel. i. p. 60 ; Keeve, Conch. Icon. pi. 77- fig. 40.5. 

Nanina nouleti, Gray, Cat. Pulm. p. 121 ; {Trochomorphci) 
Albers, Die Hel. p. GO, 2nd ed. ; Mousson, Journ. de Conch. 18G5, 
p. 190; {Xest(i) 1M70, p. Ill ; {Heiniplecta) Paetel, Cat. Conch. 
1873, p. 85; Schmeltz, Cat. Miis. Godeffroy, v. p. 90 {" Zonites" 
in error). 

Helix {Nanina) rubricata, Gould, Proc. Bost. Soc. Nat. Hist. 
1846, p. 178; Pfeiffer, Mon. Hel. i. p. 69. 

Nunina rubricata, GonlJ, Expl. Exp., Shells, p. 29, fig. GG ; Gray, 
Cat. Pulm. p. 129; {Uemijilecta) H. & A. Adams, Gen. Moll. ii. 
p. 223. 

So far as known, this species is restricted to Viti Levu and Ovalau 
Islands, where it is not uncoamion beneath decaying vegetation in 
forests. 

It may be distinguished by its large size (25 to 30 millim. in 
diam.), dark chestnut-colour, globose-turhinate form, shining sur- 
face, and chiefly by the spiral impressed striae on the body-whorl. 

2. Nanina casca. 

Helix cnlva, Gould, Proc. Bost. Soc. Nat. Hist. 18J6, p. 179; 
Pfeiffer, Mon. Hel. i. p. 41. 

Nanina ralua. Gray, Cat. Pulm. p. 129. 

Nanina casca, Gould, Expl. Exp., Shells, p. 31, fig. 69 ; H. & A. 
Adams, Gen. Moll. ii. p. 222 ; (Orobia) Albers, Die Hel. 2nd ed. 
p. 59; Mousson, Journ. de Conch. 18G5, p. 191; (Xesfa) [870, 
p. 1)2; {Thalassia) Paetel, Cat. Conch. 1873, p. 84; Schmeltz, 
Cat. Mus. Godeff. v. p. 90. 

Helix vitiensis, Pfeiffer, Proc. Zool. Soc. 1855, p. 108, pi. 32. 
fig. 9 ; Malak. Bliitt. 1857, p. 35. 

Nanina vitiensis (Xesta), Pfeiffer, Vers, p. 120. 

Eurypus cascus. Semper, Phil. LandmoU. i. p. 37, pi. 1. fig. 17. 

This species, like N. nouleti, is confined to Viti Levu and Ovalau. 
Like all the Viti species, lives beneath decaying leaves and under 
rotten wood. 

A rather solid, subconoid, more or less depressed, smooth, shining 
species, of a pale corneous colour, sometimes with a slightly ruddy 
spire and perforated base. 



I887.J MOLLUSKS OF THE VITI ISLANDS. 167 

Major diam. 15 to 21 millim. Gould's first name being preoccu- 
pied by Lowe for a Madeira Helix, he changed it to casca, 

3. Nanina pfeifferi. 

Helix j)f<^lff<'.>-i (lYanina'!), Philippi, Arcli. f. Nat. 1845, p. 62 • 
Chemnitz, 2nd ed. Helix, pi. 31. figs. 9-10; Pfeiffer, JNIon. Ilel! 
1. p. .04; Reeve, Concli. Icon. no. 1282, pi, 1F5. 

minina pfeiferi. Gray, Cat. Piilm. p. 94 ; {Xesta) Albers, Die 
Hel. p. 59 ; H. & A. Adams, Gen. Moll. ii. p. 223 ; {Xesta) 
Ifeitfer, Vers, p. 120 ; {Xesta) Mousson, Journ. de Conch. 1870. 
p. 11 1. 

Helix lurida, Gould, Proc. Bost. Soc. Nat. Hist. 184f^ d 179 • 
Pfeiffer, Mon. Hel. i. p. 4 7. 

Nanina lurida, Gould, Expl. Exp., Shells, p. 31, fig. 68 ; Gray, 
Cat. Pulm. p. 128; (Hemiplecta), H. & A. Adams, Gen. Moll. li. 
p. 223 ; {Xesta) Paetel, Cat. Conch. 1873, p. 85 ; JNIousson, Journ. 
de Conch. 1865, p. 190; Schmeltz, Cat. Mus. Godeff. v. p. 71. 

I received many examples of this species from a missionary who 
collected them at Kantavu. Dr. Griiffe obtained specimens in Viti 
Levu. 

Though I have followed Pfeiffer and Mousson in referring this 
species to pfeifferi, still I have some doubts as to their being the 
same species. Pfeiffer and Deshaves assisn it to China, and 
Cuming erroneously cites the Sandwich Island's as its habitat. 

Though nearly as large as nouleti, it may be distinguished by its 
light colour, more depressed whorls, and smooth surface. 

4. Nanina fragillima. 

Nanina {Xesta) fragillima, Mousson, Journ, de Conch. 1870 
p. 112, pi. 7. fig. 3 ; {Microcystis) Paetel, Cat. Conch. 1873, p. 8^! 

Hehx fragillima, Pfeiffer, Mon. Hel. vii. p. 91. 

I am indebted to Dr. Griiffe for examples of this species which he 
obtained in the interior of Viti Levu. It is also recorded from 
Kantavu. 

It IS a thin, transparent, shining, pale horn-coloured species with a 
depressed spire and strongly convex base. The last whorl is con- 
spicuously angulated. Major diam. 14 millim, 

5. Nanina similis, 

Euripus similis, Semper, Phil. Landmoll. i. p, 37, pi. 1. fie-. 18 
pi. 2. fig. 91. I f & , 

Nanina {Euripus) similis, Schmeltz, Cat. Mus. Godeff. v. p. 91. 
Helix similis, Pfeiffer, Mon. Hel, vii. p. 1 12. 

This species, which is uuknown to me, was collected by Dr, Graffe 
on Viti Levu. 

It is described as a solid, orbicular, depressed shell of a fuscous 
horn-colour, with irregular fuscous lines ; whorls bi, the last one 
obsoletely angulated just above the periphery, Diam. 17, height 
IO5 millim. 



168 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

6. Nanina HOYTI. 

Nanina hoyti, Garrett, Amer. Journ. Conch. 1872, p. 221, pi. 19. 
fig. 6; Schmeltz, Cat. Mus. Godeff. v. p. 91. 

Helix hoyti, Pfeififer, Mon. Hel. vii. p. 525. 

Common in Taviuni, Gomea, and Lanthala. 

It may be distinguished from N. cascu by the pale brownish 
sutural band, darker spire, tawny columella, and the coarse wrinkles 
just beneath the suture. Major diam. 19 to 20 millim. 

7 . Nanina otare.-e. 

Nanina otarece, Garrett, Amer. Journ. Conch., 1872, p. 222, 
pi. 19. fig. 8; Schmeltz, Cat. Mus. Godeff. v. p. 91. 

Helix otarece, Pfeiffer, Mon. Hel. vii. p. 524. 

I discovered numerous examples of this fine species on the north- 
west portion of Vanua Levu. 

It is about the same size as N. nouleti, and like that species has 
coarse strise of growth and faint spiral lines, the former being larger 
and much more uniform in size, and the latter on our s])ecies only 
discernible by the aid of a lens. It may be at once distinguished by 
the fulvous-brown colour and the large circular pale cream-white 
basal patch. The last whorl is also more depressed than in N, 
nouleti. Major diam. 29 millim. 

8. Nanina polita. 

Nanina nouleti, var. polita, Mousson, Journ. de Conch. 1865, 
p. 190. 

A few examples found beneath dead leaves on the eastern part of 
Viti Levu. 

It is smaller, more depressed, smoother, more polished, and the 
whorls are flatter than in N. nouleti. The base is also darker, and 
there is not the least trace of spiral striae. It may, I think, take 
specific rank. 

9. Nanina tenella. 

Nanina tenella, Garrett, Amer. Journ. Conch. 1872, p. 222, pi. 19. 
fig. 7. 

Helix tenella, Pfeiffer, Mon. Hel. vii. p. 525. 

Not uncommon in forests on the south-eastern part of the Vanua 
Levu and Kioa. 

It is a fragile, smooth, polished, transparent, whitish horn-coloured 
species shaped like N. hoyti, but only 1/ millim. in diameter. Its 
paler colour and thinner transparent texture will at once separate it 
from N. casca. 

10. Nanina godeffroyana. 

Nanina godeffro%jana, Garrett, Amer. Journ. Conch. 1872, p. 223, 
pi. 19. fig. 19. 

Helix godeffroyana, Pfeiffer, Mon. Hel. vii. p. 524. 

This fine large species was taken in considerable numbers in the 
interior of the north-east portion of Vanua Levu. 



1887.] MOLLUSKS ON THE VITI ISLANDS. 169 

It may at once be recognized by its large size (38 millim. in 
diameter), depressed turbinate form, yellowish or light brownish 
horn-colour with a wide fulvous-brown sutural band. 

11. Nanina assavaensis, n. sp. 

Shell imperforate, turbinately globose, indistinctly striated with 
rather coarse lines of growth, smooth, shining, subpellucid, smoky 
horn-colour ; whorls 5, slightly convex, the last one obscurely 
angulated just above the periphery ; aperture oblique, orbicular- 
luniform ; peristome thin, straight ; columella abbreviately reflected 
over the axis of the shell. 

Diam. 20, height 12 millim. 

Abundant beneath dead leaves at Naviti Island, one of the Assawa 
group. 

Genus Microcystis, Beck. 

1. Microcystis unisulcata. 

Nanina unisulcata, Mousson, Journ. de Conch. 1865, p. 191; 
{Microcystis) 1870, p. 113; (Thalussina) Paetel, Cat. Conch. 1873, 
p. 85 ; Schmeltz, Cat. Mus. Godeff. v. p. 90. 

Helix unisulcata, Pfeiffer, Mon. Hel. v. p. 80. 

Helix (Petasia) unisulcata, Paetel, Cat. Conch, p. 96. 

Helix laqueata, Baird, Brenchley, Cruise of the ' Curagoa,' p. 446, 
pi. xl. figs. 8, 9. 

This peculiar species is widely diffused throughout the group, and, 
like all the Yiti species, is found beneath decaying vegetation. 

It is the only species, so far as known, which exhibits a sculptured 
surface. The unisulcated whorls and more or less nodulous columella 
will readily distinguish it. 

2. Microcystis kioaensis. 

Nanina kioaensis, Garrett, Proc. Phil. Acad. Nat. Sci. 1873, 
p. 237, pi. 3. fig. 71 (" kifaensis " in error). ^/ 

Naninal taviuniensis, Liardet, Proc. Zool. Soc. 1876, p. 99, pi. 5. I 

figs. 1, la, b. 

This very distinct species only occurred to my notice in the 
mountain-forests of Kioa Island. Mr. Liardet's Nanina taviuniensis, 
which he obtained at Taviuni and Gomen, differs only from our 
species in having the base minutely perforated. 

It may be distinguished by its depressed globose form, yellowish 
horn-colour, deep rounded body-whorl, and the prominent columellar 
fold. 

3. Microcystis nodxjlata. 

Nanina (Microcystis) nodulata, Mousson, Journ. de Conch. 18/0, 
p. 1 14, pi. 7. fig. 4 ; (Thalassia) Paetel, Cat. Conch, ed. 1873, p. H5. 

Helix nodulata, Pfeiffer, Mon. Hel. vii. p. 67. 

Discovered by Dr. Graffe on Vanua Balavo, one of the Windward 
Islands. 

Its chief characters are its depressed globose form, brilliant horn- 



170 MR. A, GARRETT ON THE TERRESTRIAL [Feb. 15, 

colour, and nodulous columella. It is more depressed and paler 
than the preceding species. 

4. Microcystis excrescens. 

Nanina {Microcystis) excrescens, Mousson, Journ. de Conch. 1870, 
p. 1 15, pi. 7. fig. .'), 1871, p. 8 ; Paetel, Cat. Conch. 1873, p. 84 ; 
Schmeltz, Cat. Mus. Godeff. v. p. 91. 

Helix excrescens, PfeifFer, Mon. Hel. vii. p. G7. 

Microcystis excrescens, Garrett, Journ. Phil. Acad. Nat. Sci. 1881, 
p. 381. 

This small species was found by Dr. Griiffe on the eastern portion 
of Viti Levu and on several islands in the Tonga group. I discovered 
it on one of the Cook's Islands. 

It is nearly of tlie same size and shape as M. nodulata, with the 
columellar fold of M. kioaensis, but is much smaller than the latter 
species. 

5. Microcystis upolensis. 

Nanina upolensis, Mousson, Journ. de Conch. 1865, p. 16(J ; {Mi- 
crocystis) 18(>9, p. 327; \S7i),'VAr. oneataensis, p. 114; {Thalussia) 
Paetel, Cat. Conch. 1873, p. 85 ; Schmeltz, Cat. Mus. Godeff. v. p. 90. 

Helix upolensis, Pfeiffer, Mon. Hel. v. p. 108. 

Helicopsis vpolensis. Pease, Proc. Zool. Soc. 1871, p. 475. 

Helix samoensis, Baird, in Brenchley, Cruise of the ' Curagoa,' 
p. 447, pi. xl. figs. 12, 13. 

This species was first described from specimens collected by Dr. 
Griiffe on Upolu, one of the Samoa Islands, and I subsequently dis- 
covered it on the islands of Vanua Balavo and Oneata. 

A very smooth, highly polished, orbicular, depressed, pale horn- 
coloured species with a slight columellar nodule. 

f). Microcystis sororia. 

Helix sororia, Cox, Proc. Zool. Soc. 1870, p. 83 ; Pfeiffer, Mon. 
Hel. vii. p. GO. 

This species, which is unknown to me, was discovered by Mr. Brazier 
at Ovalau. 

A small imperforated, thin, smooth, shining species of the 
depressed-globose form, with five moderately convex whorls, of a 
uniform yellowish-olive colour, and with a simple columella. 

7. Microcystis firmostyla. 

Nanina firmostyla, Mousson, Journ. de Conch. 1865, p. 166; 
{Microcystis) 1871, p. 7; Schmeltz, Cat. Mus. Godeff. v. p. 90; 
{Microcystis) Paetel, Cat. Conch. 1873, p. 84. 

Helix firmostyla, Pfeiffer, Mon. Hel. v. p. 70. 

Helicopsis firmostyla, Pease, Proc. Zool. Soc. 1871, p. 475. 

Obtained by Dr. Gtiiffe at Tikombia, one of the Windward Islands. 
He also found it generally distributed throughout the Tonga Islands. 

A minute, depressed-convex, highly polished, horn-coloured 
species, shaped like M. upolensis, but only half as large. The colu- 
mella is sometimes slightly nodulous. 



1887.] mollusks of the viti islands. 171 

8. Microcystis perpolita. 

Nanina {Microcystis) perpolita, Mousson, Journ. de Conch. 18G9, 
p. 326, pi. 14. fig. 1; 1870, p. 113; 1871, p. 8, var. solida ; 
Schmeltz, Cat. Mus. Godeff. v. p. 90. 

Helicopnis perpolita. Pease, Proc. Zool. Soc. 1871, p. 475. 

Helix perpolita, Pleiffer, Mon. liel. vii. p. 65. 

This species was found by Dr. Giiiffe on Viti Levu. It also 
inhabits Tonga and tiie Samoa Islands. 

It is a convexly dej)ressed, pellucid, highly polished, yellowish 
horn-coloured species Avith 4^ whorls. 

9. Microcystis stearnseana, sp. nov. 

Shell small, imperforated, depressedly globose, pellucid, smooth, 
shining, luteous horn-colour ; spire convexly rounded, apex de- 
pressed ; suture linear, narrowly margined ; whorls 5, flatly convex, 
last one not descending in front, rounded, base convex ; aperture 
nearly vertical, roundly lunate ; peristome straight, acute, regularly 
curved, margins remote, not converging ; columella with a white, 
prominent, nearly horizontal tooth-like fold ; within the base of the 
aperture, a short distance from the margin, are two sublamelliform 
white teeth, the upper one the larger, crest- like, the smaller one close 
to the coluiiiellar fold. 

Major diam. 3, height 2 millim. 

Hab. Viti Islands. 

Several examples found beneath rotten wood at Vanua Balavo, 
and a few were obtained under dead leaves at Uea or Wallis 
Island. 

It is closely allied to Nanina {Gastrodonta) eiisifera, Mousson, 
a Samoan species, which is smaller, more depressed, the body-whorl 
being subangulated and more depressedly rounded. I name this 
singular species after my friend R. E. C. Stearns, Esq. 

Genus Trochonanina, Mousson. 
1. Trochonanina samoensis. 

Nanina samoensis, Mousson, Journ. de Conch. 1865, p. 165. 

Helix samoensis, Pfeiffer, Mon. Hel. v. p. 70. 

Zonites (Conula) samoensis, Mousson, Journ. de Conch. 1869, 
p.331 ; 1870, p. 116; 1871, p. 10; Paetel, Cat. Conch. 1873, p. 86 ; 
Schmeltz, Cat. Mus. Godeff. v. p. 90. 

Helix clayi, Liardet, Proc. Zool. Soc. 1876, p. 101, pi. 5. fig. 7. 

Microcystis samoensis, Garrett, Journ. Phil. Acad. Nat. Sci. 1881, 
p. 384. 

This minute species is generally diffused throughout the group. 
It is also common in the Tonga, Cook's, and Samoa Islands, and rare 
in the Marquesas. Under decaying vegetation. 

It is a perforated, thin, depressed, turbinated, reddish or brownish 
horn-coloured species, with five strongly convex whorls, the last one 
angulated on the periphery. 



172 MR. A. GARRETT CTN THE TERRESTRIAL [FfO. 15, 

2. Trochonanina MICROCONUS. 

Nanina nijcroco?i?/s, Mousson, Journ.de Conch. 1865, p. 192; 
{Thalassia) Paetel, Cat. Conch, p. 85. 

Helix microconus, Pfeiffer, Mon. Hel. v. p. 94. 

Zonites (Comtlus) microconus, Mousson, Journ. de Conch. 18/0, 
p. 117; Paetel, Cat. Coach. 1873, p. 8G ; Schmeltz, Cat. Mus. 
Godeff. V. p. 90. 

Helix pinnockii, Liardet, Proc. Zool. Soc. 18/6, p. 100, pi. 5. 
figs. 5, 5a. 

Widely diffused over the group, and, like the preceding, this species 
lives beneath decaying vegetation. It occurs also in the Tonga and 
Samoa Islands. 

A minute, perforated, conical, greyish horn-coloured species with 
5g spirally striated whorls, the last one acutely angulated and the 
base smooth. 

3. Trochonanina barkasi. 

Helix barlasi, Liardet, Proc. Zool. Soc. 1876, p. 100, pi. 5. 
fig. 6. 

This species, which is quite unknown to me, was found at Taviuni 
by >\Ir. Liardet. He describes it as follows : — 

" Shell minute, trochiform, very minutely perforated, colour 
golden horny ; whorls 5^, convex, roughly and irregularly striated, 
transversely ribbed, last whorl acutely carinated ; beneath slightly 
convex ; striae radiating from the perforation ; aperture oblique and 
triangular." 

4. Trochonanina calculosa. 

Helix calculosa, Gould, Expl. Exp., Shells, p. 48, pi. 5. fig. 63 ; 
Pfeiffer, Mon. Hel. iii. p. 41. 

Zonites (Conulus) calculosus, H. & A. Adams, Gen. Moll. ii. 
p. 116. 

Nanina calculosa. Gray, Cat. Pulm. p. 126 ; Schmeltz, Cat. Mus. 
Godeff. V. p. 91. 

Trochonanina calculosa, Garrett, Journ. Acad. Nat. Sci. Philad. 
1884, p. 22. 

A few specimens were found on the leaves of the bushes at Malolo 
Island. It is also common and widely diffused throughout the 
Society Islands. ' Numerous examples occurred to my notice at 
Dominique, one of the Marquesas Islands. 

A minute, globose- pyramidal species, witli an angulated body- 
whorl, reflexed columella, and punctiform perforation. Colour pale 
corneous ; whorls 5, convex. 

Genus Zonites, Montfort. 
1. Zonites vitiensis. 
Zonites vitiensis, Mousson, Journ. de Concli. 18G5 p. 193; 



1887.] MOLLTJSKS OF THE VITI ISLANDS. 173 

{Hyalina) 1870, p. 1 15 ; Paetel, Cat. Conch, 1873, p. 86 ; Schmeltz, 
Cat. Mus. Godeff. v. p. 90, 

Helix vitiensis, Pfeiffer, Mon. Hel. v. p. 146. 

Common beneath dead leaves in forests near the sea-shore, and 
widely distributed through the group. 

A small, umbilicated, depressed, thin, pellucid, pale horn-coloured 
species, about the size and shape of Hyalina arborea, Say, a North- 
American species. 

2. ZONITES PLICOSTBIATUS. 

Zonites plicostiiatus, Mousson, Journ. de Couch. 1870, p. 116. 

Helix plicostriata, Pfeiffer, Mon. Hel, vii. p. 197. 

Inhabits the south coast of Vitu Levu and Kantavu (Grille). 

A little smaller than the preceding species, from which it differs 
in having the body-whorl angulated, and the sculpture consists of 
fine rib-like strise and spiral impressed lines. 

3. Zonites schmeltziana, sp. nov. 

Shell umbilicated, depressed, orbicular, pellucid, shining, obscurely 
striated, pale corneous, sparsely speckled with white; spire depressedly 
convex ; suture slightly impressed ; whorls 5, nearly flat, regularly 
increasing, last one not deflected in front, depressedly rounded, upper 
portion depressed and angulated ; base convesly rounded ; umbilicus 
small; aperture obHque, orbicular-lunate ; peristome straight, sharp, 
with remote margins ; columella slightly reflected. 

Major diam. 9, height 4 millim. 

Hab. Malolo Island. 

Common beneath decaying vegetation, in forests near the sea- 
shore. 

Nearly twice the size of Z. vitiensis ; this species has the spire 
more depressed, the whorls flatter, and may at once be recognized by 
its angulated body-whorl. 

Genus Trochomorpha, Albers. 

I. Trochomorpha merzianoides. 

Helix (Trochomorpha) merzianoides, Garrett, Proc. Phil. Acad. 
Nat. Sci. 1873, p. 237, pi. 3. fig. 72. 

A rare !«pecies inhabiting Vanua Levu, where I found a few 
examples adhering to the trunks of trees. 

Its large size (22 millim.), honey-yellow base, chestnut- brown 
upper surface, which is mottled with radiating lines and spots of a 
luteons-white colour, will at once distinguish it. The upper surface 
is coloured nearly the same as T. merziaria, a Solomon-Island species ; 
but the present species may be separated by the absence of the 
basal band, darker colour, the narrower last whorl, and its narrower 
aperture. T. merziana also differs in having the upper margin of the 
peristome inflected. 



174 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

2. Trochomorpha abrochroa, Crosse. 

Heli.v abrochroa, Crosse, Journ. de Conch. 1868, p. 176; 1870, 
p. 101, pi. 1. fig. 2; Pfeitfer, Mon. Hel. vii. p. 20/. 

Trochomorpha {Discus) abrochroa, Alousson, Journ. de Conch. 
1870, p. 123, y&v. pseudoplanoi-bis. 

This species, which appears to be somewhat scarce, was found by 
me under dead wood in the mountain-forests on Kioa Island. Dr. 
Graflfe obtained it on Viti Levu. 

Easily distinguished by its thin texture, uniform luteous horn- 
colour, depressed form, sharp crowded striae, and the angle on the 
margin of the wide umbilicus. Diam. 12 miilim. 

3. Trochomorpha ludersi. 

Helix ludersi, Pfeiffer, Proc. Zool. Soc. 1855, p. 112; {Videna) 
Vers, p. 132 ; Mon. Hel. iv. p. 183. 

Trochomorpha (Discus) ludersi, Mousson, Journ. de Conch. 
1870, p. 122 (part.). 

This fine species is abundant on the trunks of trees at Gnau 
(Angau) Island. I received a few specimens from a native at 
Ovalau, where Macgillivray obtained the type examples. 

Its large size (19 miilim.), light horn-colour, and, more particularly, 
the four narrow reddish-bronn bands, two above and two in the 
base, will separate it from any other Yitian species. 

4. Trochomorpha taviuniensis. 

Helix taviuniensis, Garrett, Amer. Journ. Conch. 1872, p. 22 3 
(" Tavinniensis " typ. err.) ; Schmeltz, Cat. Mus. GodefF. v. p. 95 ; 
Pfeiffer, Mon. Hel. vii. p. 574. 

Found in numbers on the trunks of trees in damp forests on 
Taviuni. 

A little smaller and more depressed than T. ludersi, with a single 
narrow chestnut-brown submarginal band, both above and beneath, 
on a tawny-yellow ground. Out of over 300 specimens one only 
was without the bands, and was more depressed and paler than the 
type. The bands are darker and wider than in the preceding 
species. 

5. Trochomorpha tumxjlus. 

Helix iiimidus, Gould, Proc. Bost. Soc. Nat. Hist. 184C, p. 1 75 ; 
Expl. Exp., Shells, p. 62, fig. 53 ; Pfeifft-r^ Mon. Hel. i. p. 85 ; 
Mousson, Journ. de Conch. 1865, p. 194, 18/0, p. 120; Schmeltz, 
Cat. Mus. GodefF. v. p. 94. 

Nanina tumulus. Gray, Cat. Pulm. p. 128 ; {Trochomorpha) Albers, 
Die Hel. 2nd ed. p. 60 ; {Discus) Paetel, Cat. Conch, p. 85. 

This species, which is unknown to me, inhabits Viti Levu. 
Schmeltz cites Samoa as one of its habitats, which is probably a 
mistake, as Prof. Mousson does not mention it in his paper on the 
Samoan land-shells. 

It is described as a small (14 miilim.), solid, yellowish, pyramidal 



1887.] MOLLUSKS OF THE VITI ISLANDS. I/O 

shell, with a flattened base, 6-7 whorls, the last one obtuse and 
angulated at the periphery. 

6. Trochomorpha planoconus. 

Trochomorpha planoconus, Mousson, MS., Museum Godeffroy, 
1885. 

Sliell umbilicated, trochiform, rather solid, scarcely shining, 
rugosely striated ; striae rude, irregular, oblique ; colour chestnut- 
black, varigated with fulvous, gradually passing into dark chestnut- 
brown ; apex obtuse ; base dark honey-yellow, with a darker line 
near the keel ; spire elevated, conoid, with planulate outlines ; suture 
linear, narrowly margined ; whorls 7, slightly convex, slowly and 
regularly increasing, last one acutely carinated ; keel compressed and 
rugose ; umbilicus small, deep ; aperture diagonal, subrhomobidal- 
luuiform ; peristome above the keel acute and gently arched, below 
the keel thickened and concave. 

Major diam. 19, height 10 millim. 

Ono Island. 

Two examples received from the Museum Godeffroy. As com- 
pared with T. merziamides, it is much more conical, darker coloured, 
smaller, and the umbilicus is not so large. 

7. Trochomorpha fessonia. 

Helix (Trochomorpha) fessonia, Angas, Proc. Zool. Soc. 1869, 
p. 626, pi. 48. fig. 7 ; Brazier, Proc. Zool. Soc. 1871, p. 322 (part.) ; 
Schmeltz, Cat. Mus, Godeff. v. p. 94 ; Pfeiffer, Mon. Hel, vii 
p. 201. 

Trochomorpha (Discus) transarata, var. depresso-striata, Mousson, 
Journ. de Conch. 1870, p. 121. 

I received several examples of this species from Kantavu Island, 
where Mr. Brazier obtained the type specimens. On trees. Dr. 
GrJiffe found it (rare) in the interior of Viti Levu. 

Its most essential characters are its rather small size (12 millim.), 
depressed trochiform shape, crowded irregular plicate strife, brownish 
colour, and pale markings. The spire is convexly-ccnical ; whorls 6, 
with narrow white margins. Base rather flat, corneous, with a 
reddish-brown baud next to the acute white keel. Umbilicus 
small. 

8. Trochomorpha transarata. 

Helix transarata, Mousson, Journ. de Conch. 186.5, p. 194; 
Pfeifter, Mon. Hel. v. p. 183; Schmeltz, Cat, Mus. Godeff iv' 
p. 73. 

Trochomorpha (Discus) transarata, Mousson, Journ. de Conch 
1870, p. 121, pi. 7. fig. I (excl. var.). 

Helix fessonia (Trochomorpha), Brazier, Proc. Zool. Soc. 1871 
p. 322 (part.). 

When Prof. Mousson first described this species, he had only a 
single imperfect young example before him, which Dr. GrSffe 



176 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

found at Lomaloma, Vanua Balavo. Graffe subsequently discovered 
some specimens of Trocho norpha in the interior of Viti Leva, which 
Mousson (in Journ. de Conch. 18/0, p. 122) referred to his T. trans- 
arata, adding additional characters and giving a good figure of the 
same. On the same page he describes the var. depresso-striata, 
which latter is probably distinct and = T.fessonia, Angas. 

Unfortunately I lost all my Kantavu specimens of the latter 
species when shipwrecked, and have only a single dead example of T. 
fransarata from Lomaloma, which exactly coincides with Mousson's 
original description, and agrees with his figure of a Viti Levu 
shell. 

Mr. Brazier (in P. Z. S. 1871, p. 322) adds T. transarata to the 
synonymy of fessonia. Schmeltz (in Cat. Mus. Godeff. v. p. 94, 
sp. b9'iS9) quotes Mousson's var. T. depresso-striata as = fessonia, 
and, judging from the Museum numbers, he does not include the 
type, which is no. .5235 (see Cat. no. iv. p. 73), in the synonymy. 

Its small size, depressed conical form, rough irregular plicate 
striae, brownish horn-colour with interrupted radiating pale stripes, 
elevated spire, shghtly convex base, and small umbilicus are its most 
prominent characters. 

9. Trochomorpha accurata. 

Trochomorpha (Discus) accurata, Mousson, Journ. de Conch. 
1870, p. 120, pi. 7. fig. 2. 

Nanina {Microcystis) accurata, Paetel, Cat. Conch, p. 84. 

Helix accurata, Pfeiffer, Mon. Hel. vii. p. 290. 

A very rare species, found by Dr. Graife in rocks at Veria, in the 
interior of Viti Levu. 

It has the spire more elevated than T. transarata, a greenish- 
violet colour, the striae strong and irregular, the umbilicus small ; 
•whorls 7, the last oue acutely carinated with a compressed, white 
keel. Major diam. 14, height 12 millim. 

10. Trochomorpha corallina. 

Trochomorpha (Discus) ludersi, var. corallina, Mousson, Jouru. de 
Couch. 1870, p. 123. 

I found a few examples of this species on elevated coralline reefs 
in forests on Vanua Balavo, and took a large number in similar 
stations on Mango Island. Dr. Graffe found it on coral-rocks on 
Malatta and Tutuua, the latter one of the northern islands of the 
Tonga group. 

It may be described as follows : — 

Shell nmbilicated, sublenticular, solid, finely and obliquely striated, 
fulvous horn-colour, rarely whitish, with or without a marginal 
dorsal and basal dark fuscous band ; spire convexly conoid, obtuse ; 
suture depressedly margined above ; whorls 5g, slightly convex, 
slowly and regularly increasing, the last one not descending in front, 
acutely carinated ; base convex ; umbilicus small, about one seventh 
the greater diameter of the shell ; aperture diagonal, subrhomboid- 



1887.] MOLLUSKS OF THE VITl ISIANDS. 177 

lunate ; peristome simple, straight above, acute, at the base thickened 
with callus. 

Major diam. 15, height 6-7 millim. 

As compared with T. ludersi it is smaller, differs in the ground- 
colour, the umbilicus is uot so large, and the bauds are wider and 
only two in number. 

11. Trochomorpha subtrochiformis. 

Helix trochiformis, Gould (uot of Ferussac), Expl. Exp., Shells, 
p. 61. 

Helix eurydice, Moussou (not of Gould), Jouru. de Conch. 186.5, 
p. 170. 

Trochomorpha subtrochiformis, Mousson, Jouru. de Cauch. 1869, 
p. 335, pi, 4. fig. 6 ; 1870, var. albo-striata, p. 122; Pease, Proc. 
Zool. Soc. 1871, p. 474. 

Helix subtrochiformis, Schmaltz, Cat. Mus. Gode.ff. p. Qo • 
PfeifFer, Mon. Hel. vii. p. 289. 

Mousson's var. albo-striata inhabits Kanathia, one of the Wind- 
ward Islands, and the type is a Samoan species. The variety, which 
is unknown to me, is described as follows : — 

" Spira fusco-nigrescens, strigis transversis, albis, imperfectis, 
eleganter ornata, ad basin fulvescens, cum zona peripherica fusca." 
(Mousson.) 

It is probably distinct from the Samoan type. 

12. Trochomorpha themis, sp. nov. 

Trochomorpha (Discus) ludersi, Mousson (not of Pfeiffer), Jouru 
de Conch. 1870, p. 122 (part.). 

I obtained numerous specimens of this species at Vanua Balavo, 
where they were found adhering to the trunks of trees. It is also 
recorded from Oneata Island. 

Shell with a narrow umbilicus, solid, sublenticular, not shinino- ; 
striae very fine, oblique and crowded ; whitish horn-colour, with°a 
single dorsal and basal submarginal chestnut-brown line ; spire de- 
pressedly conoid, obtuse ; suture with a depressed white margin ; 
whorls 5|, depressedly convex, slowly and regularly increasing, last 
one uot deflected in front, acutely and compressedly carinated° keel 
white ; base convex ; umbilicus aljout one eighth the major diameter 
of the shell ; aperture and peristome the same as in T. corallina. 

Major diam. 14, height 6 millim. 

As compared with T. ludersi it is smaller, paler, has only half so 
many linear bauds, striae finer and the umbilicus much smaller. 
T. corallina is of a fulvous colour (very rarely whitish), the bands 
are wider and paler and the suture has not got the white margin ; it 
is also a little larger aud smoother, and lives in a different station. 

13. Trochomorpha kantavuensis, sp. nov. 
Shell widely umbilicated, thin, fragile, pellucid, depressed, lenti- 
cular, smooth, shining ; incremental striae fine, crowded, oblique ; 
Proc. Zool. Soc— 1887, No. XII. 12 



178 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 1.5, 

Inteous-corneous, the periphery margined above and beneath with a 
reddish-chestnut line, the upper one following the whorls of the 
spire ; suture with a depressed, narrow marginal line ; whorls tJ^, 
slightly convex, rather rapidly increasing, last one uot descending 
in front, acute and compressedly carinated, keel whitish ; base more 
polished than above, convex ; aperture very oblique, depressed, sub- 
rhomboid-lunate ; peristome thin, straight above, margins cou- 
verging, basal portion slightly thickened and gently arched. 

Major diam. 20, height 7 millim. 

I received several hundred examples of this very distinct species 
from a missionary who collected them in Kantavu Island. He gave 
no information in regard to its station. 

Specimens sent to the Godeffroy Museum iii Hamburg were 
referred to T. swainsonii, a Society- Island species; and examples 
sent to an English correspondent were confounded with T. metcalfei, 
PfeifPer, a Philippine species. It is probably the Helix (Fidena) 
planorbis, in " Dr. James C. Cox's Exchange List," which he 
accredits to Kantavu. The T. planorbis of Lesson was collected by 
that naturalist in New Guinea, and differs from our shell in being 
smaller, mottled with olivaceous, and in having only 5 whorls. A 
careful comparison with the description of T. metcal/eihas convinced 
me that it cannot be tlie same as the Kantavu shell. T. swainsonii is 
smaller, thicker, rougher, much more variable, and the positions of 
the lineations are different. 

Its large size, depressed form, fragile texture, luteous horn-colour 
with the chestnut-brown marginal lines, and large umbilicus will 
readily separate it from any other Vitian species. 

Genus Patula, Held. 

1. Patula iNERMis. 

Patula inerinis, Mousson, Journ. de Conch. 1870, p. 118, pi. 7' 
fig. 7. 

Helix inermis, Pfeiffer, Mon. Hel. vii. p. 167; {Patula) Paetel, 
Cat. Conch, p. 91. 

Collected by Dr. Graffe on Vanua Balavo. Beneath decaying 
vegetation. 

A small, widely umbilicated, depressed orbicular, rufous horn- 
coloured species, with fine, oblique, costulate striae and 4 slightly 
convex whorls. Diameter a trifle more than 2 millim. 

2. Patula adposita. 

Patula adposita, Mousson, Journ. de Conch. 1870, p. 119, pi. 7. 
fig. 8. 

Helix adposita, Pfeiffer, Men. Hel. vii. p. 87 ; {Patula) Paetel, 
Cat. Conch, p. 87. 

Difcovered by Dr. Graffe on Oneata Island. 

A little larger and paler than the preceding species, and the um- 
bilicus is smaller, and the acute plicate striae are uot so crowded. 



•887.] MOLLUSKS OF THE VITI ISLANDS. ] 79 

3. PaTULA PRINCEt. 

Helix princei, Liardet, Proc. Zool. Soc. 18/6, p 100 d1 5 
figs. 4, 4a. I ' i- • ' 

This species was found at Taviuni by Mr. Liardet. 

"Shell very small, depressed, with a wide and perspective um- 
bilicus; colour dark brown; whorls 3, prominently costulated • 
suture deeply impressed ; aperture round." {Liardet.) 

4. Patula irregularis, sp, nov. 

Pitys irregularis, Mousson, Mus, Godeff. 188.5. 

Shell small, umbilicated, orbicular, depressed, thin, translucent 
greyish white under a bluish horn-coloured epidermis; sculpture 
consisting of crowded, waved, raised striie, with larger ones irrec^ulirly 
intermixed; spire convex, apex flattened; suture canaliculate; 
whorls 4, convex, slowly and regularly increasing, slightly turgid 
next the suture, last one rounded, slightly depressed above the 
periphery; base convex; umbilicus deep, about one fourth the 
major diameter of the shell ; aperture slightly oblique, irregularly 
rounded ; peristome thin. ^ i o } 

Major diam. 3, height 2 millim. 

Common at Viti Levu. I received a number of specimens of 
this species from the Museum Godeffroy, Hamburg. 

A small uucoloured species closely aUied to P. rudis, a Cook's 
Island species. The irregular striae will separate it from the other 
Vitian species. 

Genus Pitys, Beck. 
1. Pitys subd^dalea. 

Patula {Endodonta) mhdcedalea, Mousson, Jouru. de Conch 1870 
p. 117, pi. 7. fig. 6. ' 

Helix subdadalea, Pfeiffer, Mun. Hel. vii. p. 258 ; {Patula) 
Paetel, Cat. Conch, p. 95. i » v / 

Pound by Dr. Graffe in Mango Island. Like all the South-Sea 

species. It IS found beneath decaying leaves and under rotten wood 

It is a little larger than Patula adposita, and of a pale horn- 

^"^1 "T 1^ ""'^^ umbilicus, planulate spire, the stri^ costulate, 

and 52 whorls. The parietal region is garnished with three spiral 

laminse, and there are four on the palate. 

Genus Placostylus, Beck. 
I. Placostylus malleatus. 

BuUmus malleatus, Jay, Revue Zool. 1842, p. 80; Gueiin Mag 
Zool. 1843, p. 61 ; Philippi, Abbild. ii. 9. p. 10, Bui. pi. 3. fig 4 ' 
Pfeiffer, Mon. Hel. ii. p. 55 ; {Charis) id. Vers, p. 1 52 ; Reeve, Conch! 
Icon pi 29. fig. 174; Deshayes, in Fer. p. 47, pi. 144. figs. 11, 12; 
Gould, Expl. Exp., Shells, p. 81, fig. 78; {Charis) Albers, Die Hel 
Isted. p. 152, 2nd, ed. p. 196; {Charis) H. & A. Adams, Gen. 
Moll. li. p. 147 ; {Charis) Chenu, xMon. Condi, i. p. 436, fig. 3201 • 



180 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

Crosse, Journ. de Conch. 1864, p. 136, 1875, p. 20; (Charis) 
Paetel, Cat. Conch. 1873, p. 98; Garrett, Amer. Journ. Conch. 
1872, p. 231. 

Charis malleatus, FrmenMd, Yerh. zool.-bot. Ges. Wien, 1869, 
p. 874. 

PlacostyJus (Charts) malleatus, Mousson, Journ. de Conch. 1870, 
p. 125 ; Schmeltz, Cat. Mus. Godeif. v. p. 92. 

This arboreal species is restricted to Viti Levu and Ovalau 
Islands. 

Its large size (55 millim.), white colour, olivaceous-brown macu- 
lations, malleated surface, oblong aperture, broadly expanded white 
lip will distinguish it. An immaculate variety is not infrequent. 

2. Placostylus fulgueatus. 

Bulimus fulguratvs, Jay, Revue Zool. 1842, p. 80 ; Gue'rin, Mag. 
Zool. 1843, p. 62; Philippi, Abbild. ii. 9. p. 10, Bui. pi. 3. fig. 2 ; 
Pfeiifer, Mon. Hel. ii. p. 55 ; (Charis) id. Vers, p. 152; Reeve, Conch. 
Icon. pi. 29. fig. 175 ; Gould, Expl. Exp., Shells, p. 80, fig. 77 ; 
(Charis) Albers, Die Hel. 1st ed. p. 152, 2nd ed. p. 196 ; (Charis) 
II. & A. Adams, Gen. Moll. ii. p. 147 ; Mousson, Journ. de Conch. 
1865, p. 195; Crosse, Juurn.de Conch. 1864, p. 137, 1875, p. 17; 
(Charis) Paetel, Cat. Conch. 1873, p. 98 ; Garrett, Amer. Journ. 
Conch. 1872, p. 231, pi. 18. fig. 1. 

Placostylus (Charis) fulgurntus, Mousson, Journ. de Conch. 1870, 
p. 125 ; Schmeltz, Cat. Mus. Godiff. v. p. 92 

Charis fulguratus, Frauenfeld, Verb, zool.-bot. Ges. Wien, 1869, 
p. 874. 

Otostomus fulguratus (Charis), Semper, Phil. Landmoll. iii. 
p. 158, pi. 17. fig. 10. 

Bulimus exiinius, Reeve, Conch. Syst. ii. p. 173. 

Flahocheilus gracilis, Broderip, Proc. Zool. Soc. 1840, p. 182. 

This, like the preceding species, is confined to the islands of Viti 
Levu and Ovalau, where it occurs on the trunks and foliage of trees. 

A rather solid, oblong, olive-yellow species, with longitudinal, 
irregular, more or less interrupted, waved olive-brown stripes. 
AVhorls 5, convex, last one with fine crowded longitudinal strise, and 
obliquely transverse anastomosing sulcations. Spire decorticated, 
whitisb or reddish. Aperture obauriform, white or light fvilvous, 
and the hp white and widely expanded. Length 45-50 millim. 

3. Placostylus elobatus. 

Bulimus elobatus, Gould, Proc. Bost. Soc. Nat. Hist. 1846, 
p. 190 ; Expl. Exp., Shells, p. 72, fig. 84 ; Pfeiffer. Mon. Hel. ii. 
p. 184 ; (Charis) id. Vers, p. 152 ; (Charis) Albers, Die Hel. 2nd ed. 
p. 196; Crosse, Journ. de Conch. 1864, p. 140 (exc\. variety); 
(Placostylus) Paetel, Cat. Conch. 1873, p. 98 ; Garrett, Amer. Journ. 
Conch. 1872, p. 232, pi. 18. fig. 2; (Euplacostylus) Crosse, Journ. 
de Conch. 1875, p. 13. 

Placostylus (Charis) elobatus, Mousson, Journ. de Conch. 1870, 
p. 124; Schmeltz, Cat. Mus. Godeif. v. j). 93. 



1887.] MOLLUSKS OF THE VITI ISLANDS. 181 

Otostomus (Placosiijlus) elobatus, Semper, Phi]. LaiidmoU iii 
p. 157, pi. 15, fig. 5. 

Bulimus colubrimis, Pfeiffer, Proc. Zool. Soc. 1860, p 138 
pi. 51. fig. 4 ; Malak. Blatt. 1861, p. 13 ; Mon. Hel. vi. p. 29 •' 
Crosse, Journ. de Conch. 1864, p. 139. 

This species is restricted to the island of Vanua Levu, where it 
occurs beneath decaying vegetation. Examples inhabiting the 
interior of the island are larger and much finer than those found in 
the forests near the sea-shore. 

My largest specimens taken in the former location are dQ millim 
ong and my smallest (adults) from near the sea-shore are only 46 iu 
length. The shape varies from oblong-ovate to elongate-ovate. It is 
solid white or ruddy beneath a fulvous epidermis, and ornamented 
with longitudinal dark green waved or zigzagged stripes, which are 
more or less interrupted. The apical whorls are usually reddish. 
Ihe aperture and hps are usually orange-red, and the throat fre- 
quently whitish. The last two whorls are minutely corrugated 
A rare variety occurs with the lips and aperture wholly white. 

4. Placostylus morosus. 

Bulimus morosus, Gould, Proc. Bost. Soc. Nat. Hist. 1846 p 190 • 
?;^P'- i^.^.P-'.^he^s. P-72, fig. 82; Pfeiffer, Mon. Hel. ii p. 56 • 
(CAam)id. Vers, p. 152 ; {Charis) Albers, Die Hel. 2nd ed pf 196 • 
(p«ns) Paetel, Cat. Conch. 1873, p. 98; Garrett, Amer. Journ! 
Conch. 18/2. p. 232 ; (Placostylus) Crosse, Journ. de Conch. 1875 
P-/0, pi. 8. fig. 1; Kobelt, Jahrb. malak. Ges. 1875, p. 225,' 

Placoshjlus (Charis) morosus, Mousson, Journ. de Conch 1870 
p. 125; Hchmeltz, Cat. Mus. Godeff. v. p. 93, ' 

Bulimus elobatus, var. minor, Crosse, Journ, de Conch, 1864 
p, 140. ' 

This very distinct arboreal species has a wider range than any 
other species inhabiting the group. It is distributed throughout 
all parts of Vanua Levu, where I collected numerous examples, I 
found It also on Rambi, Koro, Taviuni, Gomea, Lanthala, and Prof, 
Mousson, on the authority of Dr, Griiffe, records it from Viti Levu 

It IS a rather thm, oblong-ovate, uniform white, decorticated shell, 
with a large aperture and widely reflected peristome. The rough 
surface is not so conspicuously malleated as P. malleatus. There 
exists a rare abbreviated variety on the west end of Vanua Levu, 
which exhibits a few oUvaceous markings similar to those on the 
latter species. 

5. Placostylus seemanni, 

Bulimus seemanni, Dohrn, Proc. Zool, Soc, 1861, p. 207, pi 26 
fig 6; Cmsse, Journ de Conch. 1864, p, 123 ; Pfeiffei' Mon. Hel. vi! 
p. 13; Novit. Conch, lii. p. 474, pi. 102. fig. 18; Garrett, Amer. 
f°^;°-C°"ch. 18.2, p. 232; {Eumecostylus) ¥^,te\, Cat, Coneh. 
l»/3, p. 99; Crosse, Journ. de Conch. 1875, p. 10. 

Placostylus (Charis) seemanni, Mousson, Journ. de Conch. 1870, 



1S2 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

p. 126; Schmeltz, Cat. Mus. Godeff. v. p. 92 ; Canefri, Malac. 
Viagio Mag. p. 85. 

Otostomiis (Placostylus) seemanni, Semper, Phil. Landinoll. iii. 
p. 157, pi- 17. fig. 9. 

This fine large ground-species is confined to Kandavu Island, 
where it appears to be abundant. 

Though subject to considerable variation in size and shape, it 
maj-, however, be distinguished by its rather slender form, whitish 
horn-colour beneath a yellowish or olive-brown e])idermis, which is 
sometimes ornamented with waved or zigzagged stripes. The surface, 
though coarsely striated with lines of growth, is seldom marked by 
transverse corrugations. The auriculate-shaped aperture is narrow, 
white, though sometimes tinted with fulvous, and the white Up is 
considerably expanded and slightly reflected and frequently con- 
tracted above. Length from 52-77 millim. Like all the ground- 
species, it is very frequently decorticated. 

6. Placostylus kantavuensis. 

Bulimus kantavuensis, Crosse, Journ. de Conch. 1870, p. 250 ; 
1871, p. 105, pi. 5. fig. 3; 1875, p. 10 ; Pfeiffer, Mon. Hel. viii. 
p. 29. 

This is also a terrestrial species, and inhabits Kandavu Island, 
where it was discovered by Mr. Brazier. 

It is described as a rather solid, cylindrically-fusiform species, 
with longitudinal rugose striae and obsolete snbmalleations. Colour 
ruddy white, with reddish apical whorls. The epidermis is olive- 
yellow, with longitudinal waved whitish stripes. Tiie last whorl is 
subcylindrical, compressed on the middle, and the auriform aperture 
and the widely expanded peristome are wliitish. Length 43 millim. 
I have never seen an example of this species. 

7. Placostylus koroensis. 

Bvlinnis koroensis, Garrett, Amer. Journ. Conch. 1872, p. 236, 
pi. 18. fig. 9 ; Pfeiffer, Mon. Hel. viii. p. 29; {Placostylus) Crosse, 
Journ. de Conch. 1875, p. 9, pi. 1. fig. 5; Schmeltz, Journ. des 
Mus. Godeff. Heft xii. 1876, p. 161. 

Placostylus koroensis, Schmeltz, Cat. Mus. Godeff. v. p. 92. 

Occurs in great profusion on the ground in the central valleys of 
Koro Island, where it is peculiar. 

It is very closely allied to the preceding species, which it resembles 
in shape and sculpture. Colour corneous or tawny yellow, with a 
white or luteous aperture and rather widely expanded white peri- 
stome. Length 53 millim. It is very frequently distorted and very 
seldom exhibits traces of a fulvous epidermis, which is disposed in 
irregular longitudinal strips and patches. Examples sent to Mr. 
Crosse were, by that learned conchologist, regarded as distinct from 
his B. kantavuensis. 

8. Placostylus hoytf. 

Bttlimushoyfi, Garrett, Amer. Journ. Conch. 1872, p. 234, pi. IS. 



1887.] MOLLUSKS OF THK VITI ISLANDS. 183 

%. 7; Pfeiffer, Mon. Hel, viii. p. .SO; (F/acosfi/lus) Crosse, Journ. 
de Conch. 1875, p. 17, pi. 1. fig. 8. 

Placostylus hoyti, Schmeltz, Cat. Mus. Godeff. vi. p. 81. 

This beautiful terrestrial species is coufiiied to that portiou of 
Vanua Levu situated to the southward of Natawa Bay. 

Though closely related to P. elobatus in colour, markings, and 
sculpture, it is, however, readily distinguished by its abbreviated 
form, turgid body-whorl, wider aperture, expanded and reflected 
lip. Length 44 to 55 inillim. 

9. Placostylus rugatus. 

Bulimus rugatus, Garrett, Amer. Journ. Cjnch. 18/2, p. 234, 
pi. 18. fig. 1 ; Pfeiffer, Mon. Hel. viii, p. 35 ; {Placoslylus) Crosse, 
Journ. de Conch. 1875, p. 18. 

Placostylus rugatus, Schmeltz, Cat. Mus. Godeff. vi. p. 81. 

Var. CRASsiLABRis, Garrett. 

Bulimus crassilabrum, Garrett, Amer. Journ. Conch. 1872, p. 233, 
pi. 18. fig. 5; Pfeiffer, Mon. Hel. viii. p. 35 ; {Placostylus) Crosse, 
Journ. de Conch. 1875, p. 18. 

Placostylus crassilabrum, Schmeltz, Cat. Mus. Godeff. v. p. 93. 

This arboreal species is restricted to Vanua Levu, where it is 
widely distributed over the island. 

It is a rather thin, oblong-ovate species, of a whitish colour 
beneath an olive-yellow epidermis, and ornamented with small olive- 
green blotches, which are sometimes zigzagged. Length 43 millim. 
Tlie transverse rugosities do not differ from those observed on P. 
fulguratus. As compared with that species it is more abbreviated, 
the outer lip more arched and more effuse, and the aperture is more 
oblique. The base is also not so much produced. 

The variety crassilabris is more solid, and the peristome and 
parietal callus are much thicker than in the typical P. rugatus. This 
variety, of which I obtained about 200 examples, was gathered in 
the interior at a point about the middle of the length of the island. 

10, Placostylus ochrostomus. 

Bulimus ochrostoma, Garrett, Amer. Journ. Conch. 1872, p. 232, 
pi. 18. fig. 3 ; Pfeiffer, Mon. Hel. viii. p. 36 ; {Placostylus) Crosse, 
Jonrn. de Couch. 1875, p. 19. 

Placoslylus ochrostoma, Schmeltz, Cat. Mus. Godeff. vi. p. 81. 

Bulimus rumbieiisis, Garrett, Amer. Journ. Conch. 1872, p. 233, 
pi. 18. fig. 4 ; Pfeiffer, Mon. Hel. viii. p. 36 ; {Placostylus) Crosse, 
Journ. de Couch. 1875, p. 19. 

Placostylus rambiensis, Schmeltz, Cat. Mus. Godeff. vi. p. 81. 

Not uncommon on foliage at Taviuni and Rambi, rare on Gomea, 
and I obtained two examples on that part of Vanua Levu opposite 
to Rambi Island. 

It is the smallest species, so far as known, inhabiting the group. 
It is rather variable in size, ranging from 28 to 40 millim. in length, 
and its shape varies from ovate to oblong-ovate. The texture is 



184 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

lather thin, the sculpture the same as ou P. rugosus, and the colour 
ruddy corneous or whitish, often with a reddish spire. Though 
very frequently wholly decorticated, the epidermis, when present, 
is very thin, light fulvous, and usually beautifully mottled with 
green, which is occasionally disposed in zigzag pattern. The aper- 
ture and lips are more or less intense saffron-yellow, paler in the 
throat. The columellar fold is not so conspicuous, and is more 
vertical than in the preceding species. The peristome, though 
usually simple, is often slightly expanded, particularly so in the 
Rambi shells, some of which have the lip slightly reflected. 

11. PlACOSTYLUS GNATJENSIS. 

Bulimus gnauensis, Garrett, Amer. Journ. Conch. 1872, p. 235, 
pi. 18. fig. 8 (in erv. guanensis) ; Pfeiffer, Mon. Hel. viii. p. 36; 
(Placosfylus) Crosse, Journ. de Conch. 1875, p. 18. 

Placostylus gnauensis, Schmeltz, Cat. Mus. Godeff. vi. p. 81. 

This graceful species is common and peculiar to Gnau Island, 
where it lives on trees and shrubs. 

It may be recognized by its rather thin texture, oblong-ovate or 
elongate-ovate form, whitish, yellowish, or reddish horn-colour, orna- 
mented with longitudinal undulating olive-green stripes, which are 
frequently shaded off with white. The surface is rugose, with small 
transverse corrugations. The aperture is tawny yellow or reddish, 
rarely white, and the lips, which are but slightly expanded, are more 
intensely coloured than is the throat. Length 45 millim. 

12. Placostylus GRAEFFEi. 

Placostylus elobatus, Mousson (not of Gould), Journ. de Conch. 
1870, p. 124. 

Placostylus moussonii, " Griiffe," Schmeltz, Cat. Mus. Godeff. v. 
p. 93. 

Bulimus, sp., Garrett, Amer. Journ. Conch. 1872, p. 232. 

i^M^»«MswtoM5SO/i27, Crosse (not of Pfeiffer), Journ. de Conch. 1875, 

Bulimus grUffei, Crosse, /. c. p. 13 ; Pfeiffer, Mon. Hel. viii. p. 30. 

This ground-species appears to be restricted to the central portion 
of Viti Levu, where it was discovered by Dr. Griiffe. I am indebted 
to the latter gentleman for two examples of this species, which is 
about the same size and shape as P. elobatus, but is a smoother shell, 
and the colour of my two specimens is olivaceous without any mark- 
ings. The aperture is whitish, and the columellar fold is more 
horizontal than in the latter species. 

13. Placostylus vitiensis, sp. nov. 
Bulimus vitiensis, Garrett, MS. (coll. Garrett). 

Placostylus vitiensis, " Garr.," Schmeltz, Cat. Mus. Godeff. vi. 
p. 81. 

Shell umbilicated, oblong-ovate, rather solid, slightly shining ; 
rosy flesh-colour beneath a thin translucent epidermis, which is 
decorated with longitudinal olive-green zigzag stripes ; surface of 



1887.] MOLLUSKS OF THE VITI ISLANDS. 185 

tLe last two whorls with small longitudinal striae and small trans- 
verse corrugations ; spire obtuse, decorticated, minutely punctured, 
two thirds the length of the shell ; whorls 5, moderately convex^ 
the last one attenuated at the base; aperture slightly oblique, oblong, 
auriform, light fulvous ; peristome white, rather widely expanded 
and somewhat reflected ; columellar lip dilated, and the fold oblique 
aad prominent. 

Length 41, diam. 17 millim. 

I obtained 20 living examples of this species, which were collected 
by the natives at Na Viti Levu Bay, on the N.E. coast of Viti Levu. 
It is smaller and a more graceful species than F. fulguratus, and 
the base is more contracted. 

Genus Stenogyra, Shuttleworth. 

1. Stenogyra TUCKERi, 

Bulimus ^Mc^m, Pfeij9Fer, Proc. Zool. Soc. 1846, p. 30; Mon. 
Hel. ii. p. 158; {Opeas) Vers, p. 156; Reeve, Conch. Icon. pi. 68. 
sp. 481 ; (Opens) Cox, Mon. Austr. Land-Shells, p. 69, pi. 13. fig. 9 ; 
Brazier, Quart. Journ. Conch, i. p. 272. 

Stenogyra tuckeri, Albers, Die Hel. ed. 2, p. 265; (Opeas) 
Frauenfeld, Verh. zool.-bot. Wien, six. p. 873 ; Pease, Proc. Zool. 
Soc. 1871, p. 473; Garrett, Journ. Phil. Acad. Nat. Sci. 1881. 
p. 393; 1885, p. 43. 

Buliinus junceus, Gould, Proc. Bost. Soc. Nat. Hist. 1846, p. 191- 
Exp]. Exp., Shells, p. 76, fig. 87 ; Pfeiffer, Mon. Hel. ii. p! 220. 

Stenogyra juncea, Mousson, Journ. de Conch. 1869, p. 340 ; Pease 
Journ. de Conch. 1871, p. 93; Proc. Zool. Soc. 1871, p. 473* 
(Opeas) Paete!, Cat. Conch, p. 104; Schmeltz, Cat. iMus. Godeff! 
V. p. 90; Garrett, Proc. Phil. Acad. Nat. Sci. 1879, p. 19. 

Bulimus wain, Cox, Cat. Austr. Land-Shells, p. 24 ; Pfeiffer, Mon. 
Hel. vi. p. 99. 

Stenoggra upolensis, Mousson, Journ. de Conch. 1865, p 1/5 . 
(Obeliscus) Paetel, Cat. Conch. 1873, p. 104 ; Schmeltz, Cat. Mus' 
Godeff. iv. p. 29. 

Bulimus upolensis, Pfeiffer, Mon. Hel. vi. p. 100. 

Bulimus panagensis, Pfeiffer, Proc. Zool. Soc. 1846, p. 33 ; Mon. 
Hel. II. p. 156 ; (Opeas) Vers, p. 156 ; Reeve, Conch. Icon. pi. 14 
no. 76 ; (Opeas) Albers, Die Hel. p. 175. 

Subulina panayensis, H. & A. Adams, Gen. Moll. ii. p. Ill • 
Semper, Phil. Landmoll. ii. p. 137, pi. 8. fig. 15. ' 

Stenogyra panayensis, (Opeas) Albers, D°e Hel. ed. 2, p. 265- 
Martens, Ostas. Zool. ii. p. S3 (Siam), p. 376, pi. 22. fig. 8 ; (Opeas) 
Paetel, Cat. Couch, p. 104. <= ' v v y 

Bulimus diaphanus, Gassies (not of Pfeiffer), Journ. de Couch 
1859, p. 70. 

Bulimus souverbianus, Gassies, Faune Nouv. Cale'd. p. 52 nl 2 
fig. 5 ; Pfeiffer, Mon. Hel. vi. p. 98. V 'V- ^• 

Bulimus artensis, Gassies, Journ. de Conch. 1866, p. 50; Pfeiffer 
Mon. Hel. vi. p. 98. ' 

Pkoc. Zool. Soc— 1887, No. XIII. 13 



186 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

Stenogyra novettx/yrata, 'Mon^son, Journ. de Conch. 18/0, p. 126 ; 
{Subvlina) Paetel.'Cat. Conch. 1873, p. 104 ; Schmeltz, Cat. Mus. 
Godeff. V. p. 90. 

Buiimus novemgyratus, Pfeiffer, Mon, Hel. viii. p. 138. 

Stenogyra gyrata, Mousson, MS. in Mus. Godeifrcy, 1885. 

This species, which is distributed over a larger geographical area 
than any other species of land-shell, is diffused throughout all parts 
of Polynesia, the low coral-islands as well as the more elevated groups, 
and ranges throughout Melanesia, Micronesia, Australasia, the Mo- 
luccas, Philippines, Guam, Ceylon, Siam, Cochin China, China, and 
probably extends its range as far as the east coast of Africa. 

Since the publication of my paper on the Society-Ishnd land- 
shells I have received from Mr. E. L. Layard examples of Buiimus 
souverbia7ius and B. artensis, both of which are identical with 
Polynesian specimens of S. tuckeri. 

Through the kindness of Dr. Hungerford, of Hong Kong, I have 
been enabled to compare Pfeiffer's Buiimus panayensis with 
B. tuckeri, and cannot detect a single character to separate the two 
species. 

I am strongly inclined to believe that the West-Indian Stenogyra 
tsubula, Pfr., is a form of the Polynesian S. tuclcei-i, and was acci- 
dentally imported with the Tahitian bread-fruit plants nearly a 
hundred years ago. MM. Crosse and Fischtr (Journ. de Conch. 
1863, p. 361) record the West-Indian "Buiimus subula" from 
Cochin China, and give a good figure oF the same, which latter is, 
undoubtedly the ubiquitous S. tuckeri. I reproduce their remarks 
as follows : — 

" Cette espece provient de Saigon et Fuyen-Moth, ou elle a ete 
recueillie par Monsieur Michau, dans les fosses, dans la terre et sous 
les herbes. II pent sembler tres-extraordinaire de retrouver en 
Cochinchine une espece des Antilles, qui n'a guere etc signalee 
jusqu'ici qu'a Cuba, a la Jama'ique et a. Saint-Thomas. Pour ne 
conserver aucun doute a son egard, nous avons cru devoir soumettre 
un individu authentique a I'examin de M. Pfeiffer, qui a cree I'espece. 
II faut done accepter le fait, qui peut-etre, au reste, seideinent un 
accident d'acclimatation : la petitesse et la legerete de la coquille en 
question rendent cette sujjposilion vraisemblable." 

I have lately received from Ur. Hungerford several examples of 
Stenogyra, labelled " Opeas subula, Pfr., Hong Kong," which do not 
differ from the Polynesian S. tuckeri. I have several specimens of 
Stenogyra received from Dr. Gibbons, who collected them in Algoa, 
South Africa. They were labelled " Stenogyra turri/urmis, Krauss," 
but are much smaller than the latter species, and the identification is 
questionable. They are of the same size, and coincide very nearly with 
S. tuckeri. Buiimus johannius, Morelet, from the Comoro Islands, 
can scarcely be distinguished from some forms of the latter species. 

This species, which is chiefly confined to the lowlands near the 
sea-shore, is found beneath decaying vegetation and under loose 
stones. 

They vary in size, number of whorls, development of striae. 



1887.] MOLLUSKS OF THE VITl ISLANDS. 187 

convexity of the whorls, more or less open columellar chink, and in 
texture vary from thin pellucid to thick opaque cretaceous without 
lustre. The colour is whitish, pale horn-colour, sometimes with a 
light greenish tint. Animal light yellow. Length 8 to 13 millim. 

Genus Partula, Ferussac. 
1. Partula lirata. 

Partula lirata, Mousson, Journ. de Conch, 1865, p. 136. pi. 14. 
fig. 4; 1870, p. 126 ; Heynemann, Malak. BUitt. 1867, pi. i. fig. 1 
(dentition) ; Pfeiffer, Mon. Hel. vi. p. 158 ; Paetel, Cat. Conch. 1873, 
p. 104; Schmeltz, Cat. Mus. Godeff. v. p. 91 ; Ilartinan, Cat. Partula, 
p. 14 ; Obs. gen. Partula, Bidl. Mus. Comp. Zool. ix, p. 183. 

This singular Partula lives on foliage near the sea-shore. I 
obtained several hundred examples on Lanthala, and a few at Vanua, 
Balavo, and Taviuni. Dr. GriifiFe found it on Kanathia and Oneata. 

It is, so far as known, the only species of Partula with elevated 
spiral lirse. The type is pale cinereous, with the expanded lip and 
aperture white. There is a depressed white tubercle on the parietal 
wall. A tawuy-brown variety is not uncommon. 

Genus Tornatellxna, Beck. 

1. Tornatellina oblonga. 

Tornatellina oblonga. Pease, Proc. Zool. Soc. 1864, p. 673; 1871, 
p. 473; Journ. de Conch. 1871, p. 93; Preififer, Mon. Hel. vi. 
p. 264 ; Schmeltz, Cat. Mus. Godeff. v. p. 89 ; Garrett, Proc. Phil. 
Acad. Nat. Sci. 1879, p. 21 ; Journ. Phil. Acad. Nat. Sci. 1881, 
p. 398, 1885, p. 81. 

Tornatellina bacillaris, Mousson, Journ. de Conch. 1871, p. 16, 
pi. 3. fig. 5 ; Schmeltz, Cat. Mus. Godeff. v. pp. 89, 90; Pfeiffer, 
Mon. Hel. viii. p. 316. 

Inhabits all the groups from the Marquesas and Paumotus to the 
Viti Islands. On the ground in forests from near the sea-shore to 
2000 feet above sea-level. 

Its slender form, imperforate base, and nearly vertical simple 
columella will distinguish it. 

2. Tornatellina conica. 

Tornatellina conica, Mousson, Journ. de Conch. 1869, p. 342, 
pi. 14. fig. 8 ; (var. impressa), p. 16 ; Pease, Proc. Zool. Soc. 1871, 
p. 473 ; Pfeiffer, Mon. Hel. viii. p. 316 ; Garrett, Proc. Phil. Acad. 
Nat. Sci. 1879, p. 21 ; Journ. Phil. Acad. Nat. Sci. 1881, p. 399, 
1885, p. 81 ; Schmeltz, Cat. Mus. Godeff. v. p. 89. 

Cionella conica, Paetel, Cat. Conch. 1873, p. 106. 

It has the same range as the preceding species, and inhabits the 
same station. 

It is more robust and lighter-coloured thau T. oblonga, the spire 
more tapering, body-whorl larger, more compressed, parietal lamina 
more prominent, and the columella more twisted than in that species. 



188 MR. A. GARRETT ON THE TERRESTRIAL [Feb. 15, 

3. TORNATELLINA COLUMELLARIS. 

Tornatellina columelJaris, Mousson, Journ. de Conch. 1870, 
p. 120 ; Pfeiffer, Mon. Hel. viii. p. 316. 

This species was collected by Dr. Griiffe on Kanathia Island. 

It is an imperforated, elongate-conical species of a pale horn- 
colour. It differs from the two preceding species in having small 
denticles in the palate. I do not know the species, which should 
be compared with P. perplexa, Garr., and P. nitida, Pse. 

4. Tornatellina perforata. 

Lamellaria perforata, Liardet, Proc. Zool. Soc. 18/6, p. 101, 
pi. 5. figs. 8, 8 a. 

" Shell small, acute, polished, dark brown colour ; epidermis 
thin ; whorls 5|, convex, spirally striate, with a white a|)ertural 
lamina ; aperture oblique, pyriform ; columellar lip white, projecting 
from the base of the shell, expanding slightly over region of umbi- 
licus ; outer lip impressed and of a deep purple tint. 

"This shell is found embedded in the bark of dead logs. 

" Note. — The animal has the tips of the eye-pedicels bulbous. 

«' Hal. Taviuni, Fiji." {Liardet.) 

Also unknown to me. 

Genus Vertigo, Miiller. 

1 . Vertigo pedtcultjs. 

Pupa pediculus, Shuttleworth, Bern.Mitth. 1852, p. 296 ; Pfeiffer, 
Mon. Hel. iii. p. 557 ; Schmeltz, Cat. Mus. Godeff. v. p. 89 ; 
Mousson (var. samoensis), Journ. de Conch. 1865, p. 175. 

Vertigo pedicuhts, Pfeiffer, Vers, p. 177 ; (AIcbo) H. & A. Adams, 
Gen. Moll. ii. p. 172; Mousson, Journ. de Conch. 1869, p. 341 ; 
Pease, Proc. Zool. Soc. 1871, pp. 463, 474 ; Garrett, Proc. Phil, 
Acad. Nat. Sci. 1879, p. 19; Journ. Phil. Acad. Nat. Sci. 1881, 
p. 400, 1885, p. 83. 

Pupa samoensis, " MSS.," Schmeltz, Cat. Mus. Godeff. iv. p. 108 ; 
{Sphyradium) Paetel, Cat. Conch, p. 108. 

Pupa nitens. Pease, Proc. Zool. Soc. 1860, p. 439; Pfeiffer, Mon. 
Hel. vi. p. 335. 

Vertigo nitens, Pease, Proc. Zool. Soc. 1871, pp. 463, 474. 

Pupa ki/alina, " Zelebor," Pfeiffer, Mon. Hel. vi. p. 329. 

Vertigo hyaliyia. Pease, Proc. Zool. Soc. 1871, p. 474. 

Vertigo nacca, Gould, Proc. Bost. Soc. Nat. Hist. 1862, p. 280 ; 
Otia Conch, p. 237 ; Pease, Proc. Zool. Soc. 1871, p J. 463, 474. 

Pupa nacca, Pfeiffer, Mon. Hel. vi. p. 330. 

This species is common to all the Polynesian groups, and is 
generally diffused through the Viti Islands. 

Its minute size, ovate-oblong shape, hyaline texture, obtuse spire, 
rounded aperture, and the thin slightly expanded lip will readily 
distinguish it. There are usually 5 denticles iu the aperture. 



1887.] mollusca of the viti islands. 189 

2. Vertigo tantilla. 

Pupa {Vertigo) tantilla, Gould, Prnc. Bost. Soc. Nat. Hist. 1847, 
p. 197 ; Pfeitfer, Moo. Hel. iii. p. ftbl ; {Vertigo) Moussoii, Journ. 
de Coiich. 1870, p. 127 ; {Vertigo) Schmeltz, Cat. Mus. Godeff. iv. 
p. 6!) ; (Pupiiie/l/t) Paetel, Cat. Conch. 1?73, p. 10^. 

Vertigo tantilla, Gould, Ex|)l. Ex[.., Shells, p. 92, fig. 103; 
{Aiceu) IT. & A. Adams, Gen. Midi. ii. p. 1/2; Pease, Proc. Zool. 
Soc. 18/1, pp. 4G0, Aiu^, 4 74 ; Garrett, Juurn. Phil. Acad. Nat. Sci. 
1881, p. 400, 1885, p. 84. 

Pupa pleurojihuru, Shuttleworth, Bern. Mittheil. 1852, p. 296 ; 
Pfeiffer, Mon. Hel. iii. p. 560. 

Vertigo pleurophora. Pease, Prnc. Zool. Soc. 18/1, p. 474. 

Pupa dinikeri, " Zelehor," Pteiffer, iMon. Hel. vi. p. 333. 

Vertigo ihmkeri. Pease, Pioc. Zool. Soc. 1871, p. 4 74. 

Vertigo annata. Pease, Proc. Zool. Soc. 1871, pp. 461, 474. 

Pupa annata, Pleiffer, Mon. Hel. viii. p. 407. 

Vertigo (ientifera. Pease, Proc. Zool. Soc. 1871, pp. 462, 474. 

Pupa deutifera, Pleiffer, Mon. Hel. viii. p. 408. 

Ranges from the Society to the Viti Islands, This and the 
precetling species are found beneath rotten wood, under stones, and 
amongst decaying leaves. 

In shape it varies from an abbreviale-ovate to oblong-oval, and 
also in a greater or less degree in the relative proportion of the 
whorls. Ctdnur jiale corneous under a brownish, more or less 
distinctly shagreened epidermis, which in perfect examples is fur- 
nished with oblique niembiam us riblets. The last whorl, behind 
the peristome, is frequently bisulcate. 



March 1, 1887. 
Prof. W. H. Flower, LL.D., F.R.S., President, in the Chair. 

Prof. Jeffrey Bell read extracts from a communication sent him 
by Mr. Edgar Thurston, Superintendent of the Government Central 
Museum, Madras, with reference to a Batraehian of the geuus 
Cacopus. Of a specimen of C. globulosus, Mr. Thurston wrote: — 

" On laying ojien the visceral cavity, the globular shape was found 
to be due to an enormous distention of the oesophagus and stomach, 
the latter occupying nearly the whole of the abdominal cavity, and 
the remaining viscera »&.c. being compressed and lying posteriorly. 
There was no distention of the intestinal tract. The distention of 
the oesophagus and stomach was found, on section, to be caused by 
the presence in their cavities of a mass of winged White Ants {Ter- 
mites), which, when dried, weighed 326 grains." 

The colour of C. systoma during life was reported to be " prim- 
rose-yellow marbled with black, the yellow colouring-material rapidly 
dissolving in alcohol." 



Proc. Zool. Soc— 1887, No^XIV. 14 

.-.\S\\ Mi7?>v 




190 MR. SALVIN ON ORNITHOPTERA VICTORIA. [Mar. 1, 

Mr. Salvin, on behalf of Mr. F. D. Godman, exhibited a pair of 
Ornithoptera victoria;, the male of which had been hitherto unde- 
scribed. These specimens had been obtained at the end of May 
1886. by Mr. C. M. Woodford, at North-West Bay, Maleita Island, 
one of the Solomon group. The female had been known many 
years, having been described by the late G. R. Gray from a specimen 
obtained by John MacGillivray, but the locality where it was captured 
was not recorded. The hind wings of the female were more produced 
than is usual in this section of Ornithoptera ; and this had led to the 
suggestion that O. victorice might prove to be the female of 
O. tithonus, de Haan ; but tlie description which follows this note 
shows that 0. tithonus differs widely from 0. victorice, not only in 
colour but also in several remarkable points of structure. The cell 
of the primaries was very peculiarly formed, being very wide towards 
its distal end, the middle and upper discocellular nervures being very 
long ; the lower discocellular was also long, but was ranged in line 
with the sections of the median as in true Papilio ; the second and 
third sections of the median, especially the latter, were very short, 
so that the short median branches and the median itself beyond the 
cell lay very close together. The cell of the secondaries was very 
long and narrow, though normal in the female. 

Mr. Salvin read the following description of the male insect : — 

The wings are deep black ; tlie primaries, except the costa, have 
a large patch of golden green, the outer margin of which is irregular 
and ill-defined and reaches to within a quarter of an inch of the end 
of the cell ; towards the apex is a large subtriangular golden patch ; 
parallel to the inner margin and near the anal angle is an elongated 
stigma similar to that of O. priamus and its allies. The secon- 
daries, almost from the costal margin to beyond the cell, are rich 
golden green, the distal part of the cell being black, though the 
nervures closing it are green. There are also three contiguous 
submarginal golden-green spots, whereof the two nearest the anal 
angle have a large central patch of golden yellow. Beneath, the 
wings are shining golden green, with the nervures, margins, a large 
subtriangular patch over the end of the cell of the primaries, a 
series of submarginal spots at the end of each secondary nervure, 
and two lunate spots on either side of the lower radial of the prim- 
aries black. 

The antennae and prothorax are black ; the abdomen ochraceous 
grey, with a double row of spots on either side and a ventral median 
line black. 

The primaries are narrow, with hardly any perceptible anal angle, 
the outer and inner margins meeting in a continuous regular curve. 
The secondaries are elongated and narrow, and the inner margiu 
deeply incised ; the elongated hairs of the inner margin are pale 
yellow. 

Mr. Godman also sent a specimen of a male Ornithoptera 
tithonus from the island of Waigiou for comparison ; and it was at 
once obvious how very distinct this species and O. victorice were. 

Mr. Woodford, who captured these specimens, had made a large 



1887.] ON THE PROTECTIVE VALUE OF COLOUR IN INSECTS. 191 

collection of Butterflies in the Solomon and New Hebrides groups, 
the details of which it was hoped would be laid before the Society 
at a future meeting. 



The following papers were read : — 



1. The Experimental Proof of the Protective Value of 
Colour and Markings in Insects in reference to their 
Vertebrate Enemies. By E. B. Poulton, M.A., F.Z.S., 
F.L.S., of Jesus and Keble Colleges^ Oxford, Lecturer 
on Zoology and Comparative Anatomy, St. Mary^s Hos- 
pital, Paddington. 

[Eeceived February 23, 1887.] 

Introductory. — In the preparation of a short course of lectures 
which were delivered at the Royal Institution in the spring of 1886, 
I had occasion to work up the historical aspects of my subject : — 
" The Nature and Protective Use of Colour in Caterpillars." The 
results of this inquiry were thus expressed in the introductory part 
of the first lecture: — "When Darwin was investigating the bright 
colours of animals, and was elaborating his theory of their explana- 
tion as of use in courtship, he came across the brilliant colours of 
certain caterpillars and saw at once that they were a difficulty in the 
way of the theory. For caterpillars are undeveloped organisms ; 
they have been described as 'embryos leading an independent life," 
and there is no way of distinguishing the sexes by external colour or 
structure (except in a very few instances). Therefore we here meet 
with brilliant colours, often rendering the possessors conspicuous, 
which cannot be of any use in courtship. Seeing therefore that the 
bright colours must be of use in some other way, Darwin drew the 
attention of Wallace to the subject, and asked whether he could 
suggest any explanation. Wallace accordingly thought over the 
subject, and considered it as part of the wider question of the varied 
uses (other than sexual) of brilliant and startling colour, in other 
stages of insect-life and in numerous instances scattered over the 
whole animal kingdom, and he finally ventured to predict that birds 
and other enemies would be found to refuse such conspicuous 
caterpillars if offered to them. He believed, in fact, that suchlarvse 
are protected by possessing a nauseous taste or smell, and that it is to 
their advantage to become as conspicuous as possible, so that their 
enemies are warned against a repeated experience of the disagreeable 
results which follow from tasting them, that in fact the gaudy 
colouring acts as an indication of something unpleasant about its 
possessor. It was then pointed out that, as far as experiment had 
gone, it had entirely confirmed Wallace's prediction. Conversely 
Wallace argued that larvae which were inconspicuous, being coloured 

14* 



192 MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 

SO as to resemble tlieir surroundings, would be eaten when detected, 
and this prediction also seemed to receive complete confirmation." 

Tliinking over the whole line of argument and its apparently 
complete confirmation, 1 was led to anticipate that a somewhat diffe- 
rent method of conducting the experiments would lead to a modification 
and extension of Wallace's classification of the uses of colour, in the 
direction of greater elasticity- At the same time it seemed better to 
withhold the sugiicstion until 1 had taken the opportunity of sub- 
mitting it to the experimental tes-t. I was travelling in Italy a few 
weeks after delivering the lectures, and took the opportimity of cap- 
turing many individuals of a few species of South-European Lizards, 
and of one species of Tree-Fmg (JJyla arhoren, var. nieridionalis). 
I was thus able to carry out the suggested experiments, which on 
the whole yielded results which confirmed the conclusions I had 
arrived at a jjriori, and also produced other results which I had not 
anticipated. Some of these results were shortly communicated to 
the Biological Section of the British Association at Binningham 
(188t)), and an abstract is printed in the volume containing the 
papers read at that meeting. The suggested extension of Wallace's 
line of argument, which has now beenput to the proof, is as follows: — 
The acquisition of an unpleasant taste or smell, together with a 
conspicuous appearance, is so simple a mode of protection, and 
yet ex hypothesi so absolutely complete, that it seems remarkable 
that more species have not a^ ailed themselves of this means of 
defence. What can be the principle which works in antagonism to 
such a mode of protection ? For in Wallace's theory no sugjicstion of 
a true counterhalancing limit a[)peared — i. e. one which increased 
with the increasing application of this method of defence, until the 
latter received a check or, for the time being, was rendered of no 
avail, or was even turned into an absolute danger. And yet it 
seemed probable that such an antagonistic principle would appear as 
the natural outcome of the too complete success of a method of 
defence which depends on the acquisition of an unpleasant taste or 
smell together with a conspicuous appearance. If a very common 
insect, constituting the chief food of one or more Vertebrates, gained 
protection in this way, the latter animals might be forced to devour 
the disagreeable objects in order to avoid starvation. And the same 
result might be readily brought about if a scarce and hard-pressed 
form adopted the same line, and so became dom.inant, after ousting 
many species which were much eaten by Vertebrates. If once the 
Vertebrate enemies were driven to eat any such insect in spite of the 
unpleasant taste, they would almost certainly soon acquire a relish 
for what was previously disagreeable, and the insect would be in 
great danger of extermination, having in the meantime become 
conspicuous by gaining warning colours. If the reasoning be cor- 
rect, it is clear that this mode of defence is not necessarily perfect, 
and that it depends for its apparently complete success upon the 
existence of relatively abundant palatable forms : in other words, its 
employment must be strictly limited. It has, indeed, always been 
recognized that an insect may be distasteful to one Vertebrate 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 193 

enemy, but palatable to another ; and to this extent Wallace does 
point out a limit to the application of this principle of defence. But 
the counterbalancing limit which I suggested is of course entirely 
different, for I argued that a Vertebrate enemy may be forced by 
stress of hunger to eat an insect although unpalatable to it. 
Although the latter limit is thus quite distinct, it would certainly in 
time become identical with the former, as the distaste for the insect 
gradually disappeared after it had been repeatedly eaten. In fact it 
will be shown to be probable that many (if not all) of the instances 
in which an evidently distasteful insect is eaten by certain Vertebrates 
originally rose in this way. These suggested additions to Wallace's 
theory of protection by warning colours were capable of being put 
to the practical test. To achieve this object it was only necessary to 
ascertain whether an insect-eating Vertebrate could be induced by 
hunger to eat a gaily coloured and conspicuous larva which it was 
always known to refuse when other food was present, and which was 
evidently very much disliked on the few occasions of preliminary 
" tasting," which would always occur long before the time when the 
disgusting morsel would be reluctantly swallowed. I shall presently 
show that my suggestion was in every way confirmed by the test; 
but before giving an account of my own experience I will allude to 
all the previous experiments which have been made in support of 
Wallace's theory. 

I. Brightly Coloured or Conspicuous Larvce. 

At a meeting of the Entomological Society of London (see Proc. 
Ent. Snc. ser. 3, v. p. Ixxx, 1867) Wallace made his important 
suggestion as to the biological value nf conspicuous and gaudy colours 
in cater|)illMrs. It is obvious that the question of the value of such 
colours in the larval stage is almost the same as in other stages, 
and it was chiefly from the determination of the use in the latter case 
(due originally and principally to Bates) that Wallace suggested that a 
similar solution would be found to apply to the former also. Never- 
theless there are reasons why such a usethod of defence is especially 
applicable to the laival stage. I have shown that there is a spe- 
cial reason in the anatomical construction of larvae which explauis 
why these organisms require to be defended from slight injuries 
(see Trans. Ent. Soc. Lond. 188.5, pt. ii. Aug. pp. 321-323). A 
larva " may be described as a soft-walled cylindrical tube which 
owes its firirmess, and indeed the maintenance of its shape, to the 
fact that it contiiius lluid under pressure. The pressure is exerted 
by the muscular parietes of the body. The advanta;;e of this 
coristruction is as obvious as iis danger; the larva possesses a motive 
force which can be ap|ilied to any movahle part of the surface 
through the medium of thefluid.". . . " Thiscon-truction isextreniely 
dangerous ; for a slight wound entails great loss of blood, while a 
moderate injury must prove fatal. The larvae of Smerinthtis 
ocellutiis (and many others) nibble off each other's horns, and the 
wounded larvae (although they do not seem to be aware of the injury) 



194 MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 

ose a great deal of blood, and although they may recover, are 
generally stunted ; and often I am sure the loss of blood proves 
fatal. If the wound be at all extensive, the fat-body and viscera 
protrude, owing to pressure on the side distal to the wound (that 
on the proximal side having been reheved by escape of blood)." 
Therefore it is that throughout the varied means of defence possessed 
by larvse " the object is always the same — to leave the larva 
untouched, a touch being practically fatal." Wallace also originally 
expressed this peculiar danger incurred by larvse in more general 
terms, viz. " their soft and juicy bodies are so delicate that if seized 
and afterwards rejected by a bird they would almost certainly be 
killed" (see 'Contributions to the Theory of Natural Selection,' 
1875, p. 118). Other peculiar dangers of the larval stage will be 
pointed out below. Although it will be seen that brilliantly coloured 
and distasteful larvae are often tasted by their Vertebrate foes when 
impelled by hunger, all observers agree that a second trial is rendered 
less likely because of the unusual appearance which accompanies the 
unusual and unpleasant effect upon senses other than that of sight. 

Furthermore, I am now able to bring forward instances of very 
distasteful species which have no warning colours, but, on the other 
hand, are well disguised by protective tints and markings ; and a . 
comparison between the behaviour of Lizards towards these and the 
conspicuous species respectively, affords strong confirmation of the 
truth of Wallace's suggestion. It now remains to summarize the 
whole of the evidence in favour of the prediction made in 1867 ; for, 
after all, the question is purely one of evidence, and however con- 
vincingly the a priori arguments may be put, they are chiefly 
valuai)le as guides to practical investigation. And this is frdly 
recognized by Wallace, who strongly urged the practical test upoa 
the meeting at which his suggestions were first made. Experiments 
have been made by Mr. J. Jenner Weir, Mr. A. G. Butler, Prof. 
Weismann, and by myself. Jenner Weir (see Trans. Ent. Soc. Lond. 
1869, part i., April) made use of the following birds in his investi- 
gations : — Erithaca rubecula (Robin), Emberiza citrinella (Yellow- 
hammer), Emheriza schceniclus (Reed-Bunting), Pyrrhula vulgaris 
(Bullfinch), Fringilla coelebs (Chaffinch), Loxia curvirosira (Cross- 
bill), Turdus musicus (Thrush), Anthus arboreus (Tree-Pipit). He 
also used to a less extent Carduelis spinus (Siskin) and Linaria minor 
(Redpoll). 

He obtained the following results : — 

" All hairy caterpillars (experimented upon) were uniformly 
uneaten ; " viz. Arclia caja, Eriogaster lanestris, Porthesia aurijlua, 
and Orgyia antiqua. " None of these species were even examined." 
The writer believes that the hairs are not themselves disliked, but 
that they " serve as a caution to the birds that the larvae so clothed 
are uneatable." This suggestion is supported by the fact that the 
young and comparatively hairless larvse of Spilosoma meiithastri 
were tasted by the Siskin, Redpoll, and by a West-African Finch 
{Textor erythrorhynchus), but these three birds evidently found the 
larvse disagreeable, and soon left them alone. On the other hand the 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 195 

more mature larva with its characteristic warning hairs was never 
even molested. It is probable that this explanation may be true 
of this and some other species, but it obviously does not apply in 
the case of P. auriflua, &c., in which the hairs themselves are a 
source of intense irritation and annoyance. Mr. Jenner Weir found 
the same results with the spiny larvae of Vanessa urticce and F. io, 
and he draws the same conclusions as to the meaning of the spines. 

In this case the author states that " the metallic-looking chrysa- 
lides were also invariably rejected, thus showing that the spines were 
not the cause of the uneatableness of the larvae." Experiments 
were also made with the following comparatively smooth-skinned,, 
highly conspicuous caterpillars : — Abraxas grossulariata, Diloha ccb- 
ruleocephala, Anthrocera JilipendultB, and Cueullia verbasci. In no 
case were these species molested. Thus these experiments strongly 
confirm Wallace's prediction. It may be doubted whether the 
larvae of Arctia caja and of Spilosoma menthastri can be included 
among the brightly-coloured larvae intended by Wallace, but there 
is no doubt that the habits of these species are such as to reader 
them conspicuous in spite of their sober coloration. In Trans. Ent. 
Soc. Loud. 1870 (part iii., August), Jenner Weir has contributed 
another paper on the same subject. Mr. H. D'Orville, in the 
'Entomologist's Monthly Magazine' (vol. vi. p. 16), had affirmed 
that the larvae of Cueullia verbasci are eaten by birds in the wild 
state. In his second paper Jenner Weir conclusively showed that 
this species was not eaten in the wild state in certain localities, and 
he again proved that it was not touched in his aviary. It seems 
therefore certain that Jenner Weir is correct as far as his species 
of birds are concerned ; but at the same time D'Orville seems to 
prove that this distasteful species may be eaten by certain birds. In 
this paper Jeuner Weir confirms his previous experience with regard 
to E. lanestris, D. cceruleocephala, A. grossulariata, and P. auriflua. 
He also includes the following new species in the list of gaudy or 
conspicuous larvae which were untouched by the above-mentioned 
birds: — Orlonestis Rotatoria, Lasiocampa quercus, CUsiocampa 
neustria, Hybernia defoliaria. Of these the two first are hairy, and 
although with sober colour, are generally conspicuously placed on 
their food-plants. (I think it is also exceedingly probable that their 
rejection may be partially due to the possession of irritating hairs.) 
The two last-named larvae are certainly brightly coloured. 

Mr. A. G. Butler (Trans. Ent. Soc. Lond., March 1869, p. 27) 
only records experiments with three species of conspicuous larvse. 
Lacerta viridis always refused the larva of A. grossulariata, but 
devoured that of Phragmatobia fuliginosa. The latter is not 
brightly coloured but, like the larva of S. menthastri, it is hairy and 
not inconspicuous. Frogs also refused the conspicuous larvse of 
A. grossulariata and Halia ivavaria, although this was often after 
tasting them, the rejection being accompanied by evident signs of 
disgust. The former larva was also rejected by Spiders, either with 
or without preliminary seizure. It is noteworthy that the larvae 
would certainly be uninjured after being seized and then relinquished 



196 MR. E. B. POULTON ON THE PROTECTIVE [Mat. 1, 

by the Frogs, and probably in the case of the Spiders also (in fact, 
Mr. Butler states that this was the case), but a bite from a Lizard 
would always be very serious and generally fatal. Mr. Butler states 
that the Lizards seized the distasteful larvae before rejecting them, 
although this may not have been in many instances. Mr. Butler 
has kindly given me an account of some further experiments upon 
birds, the results of which are included in the Tables below. 

Professor Weismann (' Studies in the Theory of Descent,' PartIL, 
pp. 336-340, English translation by Prof. Meldola) also experimented 
Vi\i\\ Lacertu viridis, with the results tiiat the fi)llowing brightly 
coloured distasteful larvae were refused : — C/isiocnmpa neustria, 
Euchelia jacobcEcE, Pyycera bucephala, Pieris brnssiccB, Deilephila 
gain. On the other hand, the very highly conspicuDus larvae of 
Deilephila evphorhitB were eaten, as also were those of E. lanestris 
and Lusiocampa pini. The young larvae of Lasiocumpn rubi, at a 
stage when they much resembled those of the rejected E. jacobeeee, 
were eaten after cautious examination. Professor Weis.nann also 
regarded the larva of Pnpilio mavlinon (always rejected) as conspi- 
cuous ; but from my own experience I should certaiidy consider it well 
concealed upon its food-plant, and I sliould exchide it from the cate- 
gory of conspicuous larvae which support Wallace's sugj:e»tion. 

Mv own experiments were conducted with green Tree-Frogs {Hyla 
arborea, var. merif/innalis), and with Lizards of tlie follownisj spe- 
cies: — Laceitn viridis, L. murulis (chiefly var. tiliyuerta), and Taren- 
tola maiiritanica. 

My diary, printed in the form of Appendix II. to this paper, will 
give all the details, dates, &c. of the various experiments made 
during the summer of IbSG, so that it is unnecessary to further 
allude to them here. 

Finally, wlien a few weeks ago I told Mr. Jenner Weir of my 
intention to bring toiretlirr all the experimental evidence upon this 
subject, he most kindly sent me the notes of his own observations 
dnrmg lh8(j, for incorporation in this paper. His experiineiits were 
conducted wi'li the following species ol Lizards: — Ldcerta viridis, 
L. Off His, and Zootoca vivipara, and the diary is also piinted ia full 
as Appendix 1. 

It now remains to tabulate the results of all the experimental 
evidence upon conspicuous larvae detailed above or desci led in tiie 
Appendices. Before doing so, it is necessary to recall Wallace's 
original suggestion, " that brilliant or conspicuous larvae wcmld be 
found to be refused by thc-ir enemies: " that is to say, they wdl be 
found to possess some unpleasant attribute. This may be a dis- 
agreeal*le taste or a nauseous smell in the fluids and tissues of the 
larva, and perceived alter it has been bitten ; or it may be a strongly 
smelling fluid, discharged by certain s|)eciiil glands on the approach 
of an enemy (e. g. Portkesia aurijlua witli dorsal glands, or the 
Ilvnienopterons Croesus septentrioiudis with ventral glands ; in both 
these cases the smell given off from the everted glands can be readily 
perceived as sbar|) or unpleasant to ourselves). The iarvpe of Pieris 
brassica, or of Pygcera bucephala, &c., form instances of the former 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 197 

class, although one cannot he sure that there is not some smell, 
given off from the general surface of the body. It will be shown that 
in some cases it is even likely that larvae may be protected by their 
reputation for being indigestible. Again, the larvae may be disliked 
because of the possession of irritating hairs, as in the case of Porthesia 
aurijiua, in which the effects of tlie hairs are almost immediate and 
intensely irritating (to man, and evidently to lizards &c.) ; or as in 
other hairy larvae which cause irritation after longer contact (e. g. 
Odonestis potntoria, from my own experience after long handling, 
and, as I hear from others, with Lasiocampa rubi and L. quercus 
&c.) ; but there is no doubt that the effects upon the delicate skin of 
the mouth would be much more rapid in all cases. We also see that 
more than one unpleasant attribute may be present in a single larva, 
as in the case of P. aurijiua, &c. 

Just as there may be many ways in which a larva may be un- 
pleasant to its foes, there are many ways by which it may be rendered 
conspicuous, some of which have been suggested since Wallace's 
original hypothesis. Thus a larva may be conspicuous from its 
startling coloration (e. g. P. avrijlun or A. grossiilariata), or because 
it freely exf>oses itself, while its colours, although sober, do not har- 
monize with its food-plant (e. g. O. potntoria on grass, or B. rubi ou 
heather ). Again, it may become conspicuous by living in companies, 
in whuh case the individuals n^ay be brightly coloured (e. g. C. neu- 
striii, E. jacobee(e, P. bucephala, &c.) ; or may be sober-coloured, 
but stronglv contrasted v\iili the food-plant (e.g. the dark larvae of 
Vanessa io or V. urticee, freely exposed in companies on the tops of 
nettles). It is obviously of less importance for the gregarious species 
to be as conspicuou.'-lv coloured as the isolated larvae, because the 
nunil)ers add greatly to the efficiency of coni|)aratively sober colours. 
This exf)lanation ot the use of the gregarious habit in many species 
was made by Fritz Midler in ' Kosnios,' Dec. 1877, and an abstract 
of the paper was coniinunicated to the Entomological Society of 
London by Professor Meldola (see Proc. 1^78, p[). vi & vii). The 
descii[)tions of appearance in all the tables fre principally taken 
from Newman's works, the habits being cliiefly descril)ed from my 
own experience. Prof. Westwood has most kiiidlv assisted me in the 
search tor the names of many of the species employed in the experi- 
ments. (See Table I., pp. l'<J^-203.) 

A second small group of larvae must he tabulated separately, i. e. 
those which take advantage of two methods of protection which at 
first sight appear to be mutuallv exclusivt — the Tiiethod of protective 
resemblance and that of a conspicuous appearance, warning of 
un|jlc-asant attributes. Such larvae are apt to pass unnoticed because 
of the harmony between their colours and markings and the artistic 
effect of their surroundings ; but it discovered, or even if an enemy 
approach so that there is danger of their being discovered, the 
protective attitude is instantly changed for one which renders the 
larva conspicuous and warns the enemy of the presence of unpleasant 
attributes (taste or smell), or alarms it by the resemblance of the 
new appearance to some object of terror. These facts may even be 



198 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table I. — Undoubtedly 



Species 

of 
Larva. 


Method by which rendered 
conspicuous. 


Unpleasant 
attribute. 


Results of 


J. Jenner Weir, 

using many species 

of Birds and Lizards. 


A. G. Butler, 

using Birds, Lacerta 

viridis, Frogs, and 

Spiders. 


Pieris hras- 

siC(B. 


Chief colours yellow and blu- 
ish green with black spots ; 
also gregarious and freely 
exposed on upper sides of 
cabbage-leaves &c. 


Taste or smell. " Dis- 
agreeable odour 
when crushed " 
( Weismann). 










Vanessa 
io. 


Intensely black, with minute 
white points ; bristles ; also 
gr?garious and freely ex- 
posed on upper sides of 
nettle-leaves &c. 


Ejects a green fluid 
from mouth when 
touched. ? Taste or 
smell. Proof lies in 
what follows. 


Disregarded by all 
the birds. 






Vanessa 
urtica. 


Same as V. io in aU respects 
except colour, which is 
lighter, although much 
darker than leaves of net- 
tle ; yellow often present 
on dark ground-colour. 


Ejects a green fluid 
from mouth when 
touched. ? Taste or 
smell. Proof lies in 
what follows. 


Disregarded by all 
the birds. 




1 


Anthroeera 
filipen- 
dultB. 


Yellow and black ; conspicu- 
ous position on trefoil &c. ; 
so abundant locally as to 
be almost gregarious. 


? Taste or smell. Proof 
lies in what follows. 


Disregarded by all 
the birds. 






1 


Beilepkila 
euphorbia. 


Black, red, and yellow or 
white; most conspicuously 
coloured and freely ex- 
posed on the spm-ge. 


" If interrupted they 
spit out a quantity 
of green liquid of an 
acid and disagree- 
able smell, similar to 
spurge-milk, only 
worse '' {Melhtdsh, in 
Stainton's 'Manual'). 








1 


Beilephila 
gala. 


Very variable colours, but 
always strongly contrasted, 
and "almost as conspicuous 
SlS D. euphorbia; rests fully 
exposed by day on the 
stem" [of Galiuni] (Weis- 
mann). 


? Taste or smell. Proof 
hes in what follows. 








i 



1887.] 



VALUE OF COLOUR AND MARKINGS IN INSECTS. 



199 



Conspicuous LarvcB. 



Experiments. 


How far support given to 
Wallace's suggestion, 
. that brilliant and conspicu- 
ous larvae woidd be 
refused by some at least 
of their enemies. 


How far support given to 

Poulton's suggestion, 

that a limit to the success of 

this method of defence 

would result from the 

hunger which the success 

itself tends to produce. 




A. Weismaiin, 

using 
Lacerta viridis. 


E. B. Poulton, 

using three species 

of Lizards smd 

Hyla. 




Refused by L. vi- 
ridis. 




Strong support. 


No evidence, for other food was 
not withheld. 












Strong support. 


No evidence, for other food was 
not withheld. 












Eaten freely by La- 
certa muralis. Not 
offered to others. 


Support from behaviour of 
birds ; shows that a larva 
may be disliked by one insect- 
eating Vertebrate and not by 
another. 


No evidence, as above, from 
birds ; of course the sugges- 
tion cannot apply to Lacerta 
muralis, which eat the larva 
freely. 












Strong support. 


No evidence. 










Eaten at once by 
L. viridis. 




A difficulty, especially as also 
" sea-gulls and terns devour 
them in numbers " (^New- 
man). 


The correlation of a startling 
appearance with some unplea- 
sant attribute must probably 
have existed once if not now. 
Have we a case in which hunger 
oropportunity have caused the 
enemies to neglect the latter, 
and therefore to benefit by 
the former ? 








Neither examined 
nor touched by 
L. viridis. 




Strong support. 


No evidence. 







200 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table I. 



Species 

of 
Larva. 



Diloha C(B- 
rideoce- 
phala. 



Fygara bit- 
cephala. 



Orgi/ia an- 
tlqva. 



Porthena 
auriflua. 



Euchi lia 
jacob(B(B. 



Lasioeampa 
(Demlroii- 
mus) pini 



Metbod by wbieh rendered 
conspicuous. 



Yellow, green, and black ; 
freely exposed on leaves of 
hawthorn &c. 



Yellow, orange, and black ; 
downy ; gregarious ; most 
conspicuous on oak, elm, 
lime, birch, &c. 



Blnck and pink, with hairy 
tufts; freely exposed on 
upper sides of leaves of 
nearly all garden plants. 



Unpleasant 
attribute. 



? Taste or smell. Proof 
lies in what follows. 



? Taste or smell. Proof 
lies in what follows. 



Eversible dorsal glands 
doubtless yielding 
odoriferous secre- 
tion. Hairs also ap- 
parent ly disliked , and 
perhaps irritating. 



Black, red, and white ; bairj- ; Eversible dorsal glands 



very conspicuous on upper 
sides of leaves of hawthorn 
&c. 



secretion volatile and 
irritant. Hairs in- 
tensely irritating. 



Alternate rings of black and 
yellow ; gregarious ; very 
conspicuous on ragwort. 



? Taste or smell. Proof 
lies iu what follows. 



" Variegated with red, brown, 
grey, and white, with two 
blue fasiijc npar the head, 
spotted at tiie sides with 
red; it is tufted with hairs, 
one thicker than the rest 
near the tail." Apparently 
conspicuous on its food- 
plant — pine. ( ]\'esht'oodand 
Humphreys, 'British Moihs.') 



Curtis states that 
Walker found the 
hairs intensely irri- 
tating on handling 
the larva. 



Eesults of 



J. Jenner Weir, 

using many species 

of Birds and Lizards. 



Disregarded by all 
the birds. Expe- 
riment repeated a 
second season, 
when the larva 
was " examined 
when moving, but 
not eaten." 



Eaten by Lacerfa 
ar/ilis, but evi- 
dently disliked 
and generally 

avoided. 



Disregarded by all 
the birds. 



Disregarded by all 
the birds. Expe- 
riment repeated a 
second season. 
Refused by all the 
Lizards. 



A. G. Butler, 

using Birds, Laceria 

viridis, Frogs, and 

Sj)iders. 



A young Missel- 
Thrush reared 
from the nest has 
frequently eaten 
the larva', but th 
long hairs were 
always rubbed off 
before swallowing. 



Eaten, without hesi- 
tation, by a young 
Sky-Lark, which, 
however, died soon 
aCterwards with 
syniptums which 
may ha\e been due 
to irritation from 
the hairs. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 



201 



{continued). 



Experiments. 



A. Weismann, 

using 
Lacerfa viridis. 



Avoided 
ridis. 



by L. 



E. B. Poulton, 

using three species 

of Lizards and 

Hi/la. 



Eaten by Tery hun- 
gry L. miiralis, 
and, I believe, by 
L. viridis, and yet 
evidently disliked 
by all. 



How far support given to 
Wallace's suggestion, 
that brilliant and conspicu- 
ous liir\£e would be 
refused by some at least 
of their enemies. 



Strong support. 



Strong support. 



How far support given to 

Poultons suggestion, 

that a limit to the success of 

this method of defence 

would result from the 

hunger which the success 

itself tends to produce. 



No evidence. 



Strong support. 



Strong support in JennerWeirs 
observation ; and Butler's 
shows that the hairs are much 
disliked. 



No evidence, for the Missel- 
Thrush appeared to relish the 
larvae. 



L. inuralis, when 
hungry, bit the 
larva, retaining it 
for a long time, 
but in the end re- 
jected it, and much 
irritated by hairs. 



Strong support, on the whole. 
It is impossible to decide 
whether the Sky-Lark was 
killed by the larvtB. If so, it 
strongly opposes the theory of 
any instinctive knowledge. 



It is certainly a support to the 
suggestion that a Lizard when 
hungry enough should make 
such a determined attempt to 
eat the larva, which it evi- 
dently disliked. 



Entirely disregard- 
ed by L. viridis 
until after another 
similar but pala- 
table larva had 
been introduced ; 
then tasted, but 
rejected. 



Seized and relin- 
quished by hun- 
gry L. muralis, 
Probably eaten 
later ; but insuf- 
ficient evidence. 



Strong support. 



It is certainly a support to the 
suggestion that a Lizard when 
hungry enough should make 
such a determined attempt to 
eat the larva. 



Devoured by L. vi- 
ridis, "but not 
exactly relished." 



Support. 



Strong support. Eaten, al- 
though unpleasant in some 
way. 



202 



MR, 



E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table I. 



Species 

of 
Larra. 


Method by which rendered 
conspicuous. 


Unpleasant 
attribute. 


Eesults of 


Jeuner Weir, 

using many species 

of Birds and Lizards. 


A. G. Butler, 

using Birds, Lacerta 

viridis, Frogs, and 

Spiders. 


Eriogastcr 
lanestris. 


Black, red, and white ; gre- 
garious ; living on a web ; 
rather hairy ; very conspi- 
cuous on hawthorn. 


? Taste or smell. Proof 
lies in what follows. 
The hairs may be 
irritating. 


Disregarded by all 
the birds. Expe- 
riment repeated a 
second season. 






Clisiocampa 
neustria. 


Orange-red, black, wlute, and 
blue ; rather hairy ; gre- 
garious, and living on a 
web when young ; very con- 
spicuous on apple &c. 


? Taste or smell. Proof 
lies in what follows. 


Disregarded by all 
the birds, al- 
though very hun- 
gry. Eaten by L. 
viridis and L. agi- 
lis, although some- 
times refused, and 
evidently disliked. 






CueuUia 
verbasci. 


Green, yellow, and black ; 
gregarious and very con- 
spicuous on upper sides of 
leaves and on the stem of 
mullein. 


The larvae eject a 
green fluid from 
their mouths when 
disturbed. ? Taste 
or smell. Proof lies 
in what follows. 


Disregarded by all 
the birds. Expe- 
riment repeated a 
second season. 






Halia wa- 
varia. 


Green to lead-colour, with 
yellow and black. Does not 
assume the characteristic 
highly protective attitude 
so common in Geometrae ; 
but most conspicuous on 
currant and gooseberry. 


?Taste or smell. Proof 
lies in what follows. 




Always refused by 
Frogs after tasting 
them ; so also 
with Spiders. Sup- 
plied to the young 
of the Great Tit by 
the parent birds, 
and always eaten 
greedily. 




Abraxas 
grossida- 
riata. 


Cream colour, black, and 
oi-ange : as above, unlike 
most Geometrse, but most 
conspicuous on blackthorn, 
gooseberry, &c. 


? Taste or smell. Proof 
lies in what follows. 


Disregarded by all 
the birds. Expe- 
riment repeated a 
second season. 
Once eaten by L. 
agilis ; often tas- 
ted and refused; 
evidently much 
disliked. 


Always refused by 
Frogs and L. viri- 
dis after tasting 
them; so also with 
Spiders. (Epeira 
diadema and Ly- 
cosa ?, sp., were the 
Spiders used in the 
case of this and the 
previous species.) 


Hybernia 
defoUaria 


Brown and yellow ; as above, 
unlike most Geometrre, but 
conspicuous and oft<>n hang- 
ing by a thread from its 
food-plant (oak &c.). 


? Taste or smell. Proof 
lies in what follows. 


Disregarded by aU 
the birds. 







1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 



203 



(continued). 




204 MR. E. B. POULTON ON THE PROTKCTIVE [Mar. 1, 

true of gregarious larvae. Thus a group of phvtojihagous Hymeno- 
plerous larv;e may remiiiu inconspicuous while un iisturbed, but 
nevertheless the approach of an eneiny determines united movements 
iu the colony vvliich render tlie whole strikingly conspicuous, and which 
may be attended later by the emission of au offensive smell from the 
numerous ventral glands of all the individuals sinniitaneonsly {e.g. 
Croesus septentrionalis). In the otlier larvae which suddenly assume 
a terrifying attitude " the effects produced approximate somewhat to 
au intensely exaggerated caricature of a sort of jjeneralized vertebrate 
appearance, prohaljiy of the serpent type (at any rate in ClicBrocampci), 
such as would be most efficacious in the case of birds. It is likely 
that the terrifying appearance of our own larvae in temperate 
latitudes first arose in the tropics, where the imitated cause of alarm 
to the enemies of the larva is real and obvious. And it is prubable 
that the success of the same method in countries where the re|)tiliaa 
fauna cannot be said to constitute a source of alarm is due to 
inherited memories of a tropical life which live on, as that instinctive 
fear of anything snake-like which is so commonly exliibited by the 
higher land-vei tebrates including ourselves." (Poulton, Trans. £nt. 
Soc. Lond. 1886, pt. ii. June, pp. 15(), 157). 

The success of this combination of defensive measures depends on 
the extraordinary sensitiveness of the larvae, so that the transition 
from the one method to the other is instantaneous, and in the CMse 
of the suddenly assumed terrifying attitudes, the enemy is additionally 
alarmed by the way iu which some dreaded ohject seems, as it were, 
to spring into existence. It is very unfortunate that so few expe- 
riments have beeu made upon this most interesting group of 
larvae. 

Just as it was suggested that insect-eating Vertebrates might, 
under the influence of huni;er, be induced to eat and finally to relish 
distasteful larvae, so we must expect that the same cause would in the 
end prevent this elaborate system of intimidation from being success- 
ful. In this case, however, there is no prejudice against an unplea- 
sant taste or smell to be overcome, and it is most probable that the 
larvae would be in great danger as soon as the imposition was detected. 
It is perhaps on this account that these methods are adopted by an 
exceedingly small proportion of larvae, but also because a certain size 
is necessary for any chance of success. Nevertheless this size is 
less than might be anticipated, for the anterior part of the body with 
large eye-like marks is generally swollen out into a resemblance to 
the head of a serpent, while the larval body is partially concealed 
among the leaves of the food-plant, and, in many positions, what is 
seen merely serves to suggest a far more extended length than that 
which actually exists. Wallace has suggested that it is very probable 
that the " spectacles " of the Cobra are terrifying marks, which warn 
the enemy against approach, and it is most interesting to note that 
the CA<5Broca»«jpa-larvae mimic the terrifying eye-like marks of a Cobra- 
like serpent, and not the real eyes of a serpent, which are relatively 
small. (Table II., pp. 206, 207). 

Having thus tabulated the results of experiments upon undoubtedly 



J887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 205 

conspicuous larvae, in every way typical of the strongly coloured 
group to which Darwin had called the attention of Wallace, and 
having further tabulated those which become conspicuous on the 
approach of danger, it is now necessary to add a few other species 
which cannot be regarded as typical of the above-mentioned class, 
but which are not concealed or are very imperfectly concealed by 
protective colouring, which are more or less freely exposed upon their 
food-plants, or about which a difference of opinion exists. (Table 
III., pp. 208, 209.) 

We will now consider a few of the conclusions arrived at by a study 
of the above tables, which give the whole of the experimental evidence 
(as far as I am aware) upon the precise question originally raised by 
Darwin. The first and obvious result of the first table is, with only 
one entirely antagonistic exception, the most complete demonstra- 
tion of the truth of Wallace's suggestion, that a highly conspicuous 
appearance would be found to be accompanied by some unpleasant 
attribute. The exception is very remarkable, as the larva is so 
highly coloured, and I think the total results of all the experiments 
will justify us in concluding that the larva of jD. euphorbice 
is unpleasant to some as yet unknown foes, and in all probability that 
it has been recently distasteful to a larger number. As to the 
results which bear upon my own suggestion, it must be observed 
that the only considerable support is to be expected from the 
columns of experiments under my name, because the other observers 
did not enter upon the investigation with this object in view, and 
therefore did not test whether a distasteful form would be eaten 
when other food was withheld. It will, however, be found that 
when this test was applied, in nearly all cases the unpleasant larv« 
were either swallowed, or a most determined attempt was made to 
eat them. And there is some incidental support in the other 
experiments also; for in many instances the larvoe were " tasted " 
before being rejected, aud in other cases even stronger con- 
firmation is forthcoming, when the iarvse were eaten, although 
" not exactly relished " (Weismann). Since the above was written, 
Jenner Weir's experiments in 1886 have been included, and these 
strongly confirm my own observations. 

It may be taken as proved that the continued si)read of some dis- 
tasteful form and the corresponding diminution in edible species would 
lead to the former becoming the prey of insect-eating Vertebrates ; 
for a point would ultimately be reached, as it was reached in many 
of my experiments, when hunger would become a stronger stimulus 
than those lesser prejudices in which a species can very well afford 
to indulge while palatable food is abundant. This prejudice against 
peculiarities in taste having been overcome in confinement, there is 
nothing in the conditions of natural life which could prevent the 
same result from being reached, as doubtless it has been reached, 
again and again. A comparison of all experiments of this kind ever 
made with insects will show that the likes and dislikes of insect- 
eaters are purely relative, and are manifested to a marked extent 
when they are offered a variety of insects, even when obviously 

Proc. Zool. Soc— 1887, No. XV. 15 



206 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table II. — Larvce which only become 



Species 

of 
Larva. 


Method by which 
rendered 

conspicuous. 


Unpleasant attri- 
bute real or 
imaginary. 


Eesuits of 


A. Weismann. 


Lady Verney. 


Chcerocarn- 
pa elpe- 
nor. 


Larva brown, sometimes 
green. When approached, 
the anterior part of the 
body is distended, and re- 
sembles a serpent-like head 
(of the Cobra type), while 
the real head is drawn in. 
Two pairs of large eye-like 
marks are seen by an enemy 
approaching from above or 
the side, while another pair 
meet an enemy coming Irom 
the front, and these last are 
modified in the terrifying 
attitude out of other mark- 
ings. (Poulton, Trans. Ent. 
Soc. Lond. 1886, pt. ii. 
p. 154 &c.) 


Suggested danger. 


A tame Jay ate the 
larva at once ; 
Sparrows and 
Chaffinches (wild) 
were frightened 
by it, and would 
not come near the 
seed - trough in 
which it was 
placed ; Fowls 
were evidently 
frightened, but in 
the end cautiously 
attacked it, when 
it was soon eaten. 


Kefused by small 
birds, which were 
evidently fright- 
ened by it, and 
would not come 
near a tray with 
crumbs on it, on 
which the larva 
had been placed. 
('Good Words,' 
1877, p. 838.) 

1 


Cerura vi- 
mda. 


Green and purplish brown. 
When attacked, the head is 
drawn in and a bright red 
margin inflated, with two 
intensely black marks on it 
in the appropriate position 
for eyes ; this terrifying 
face is turned towards any 
point at which the larva is 
touched ; also pink whips 
are swiftly protruded from 
the two prongs in which 
the body terminates. 


Terrifying aspect ; the 
whips might be 
thought to be stings ; 
the larva also ejects 
strong formic acid 
to a considerable 
distance. (Poulton, 
Trans. Ent. Soc. 
Lond. 1886, pt. ii. 
p. 167.) 


Devoured by L. vi- 
ridis. 


1 


- 


The phyto- 
phagous 
hymeno- 
pterous 
Crcesiis 
septentri- 
onalis. 


Green, orange, and black. 
Gregarious ; but not conspi- 
cuous until appro.iched ; 
then most conspicuous, all 
individuals whipping about 
with the abdomen and 
everting the ventral glands. 


Many median ventral 
glands everted on 
approach of enemy, 
and producing a 
most unpleasant 
smell. 




j 




A phyto- 
phagous 
hynieno- 
pterous 
larva, 
probably 
Fematus 
ribesii. 

i 


Green, with yellow and black 
markings. Gregarious, be- 
coming conspicuous when 
approached, as in the last 
species, but without ventral 
glands. 


? Taste or smell. Proof 
lies in what follows. 

1 








I 
1 
1 

1 
I 



1887.] 



VALUK OF COLOUR AND MARKINGS IN INSECTS. 



207 



conspicuous when approached and detected. 



Experiments. 



J. Jenner Weir. 



E. B. Poulton. 



Refused by Lizards. 



Eaten at one time by 
nearly all Lizards 
fi'eely ; at another 
seized without sus- 
picion, but aban- 
doned by L. mii- 
ralis, although a 
very hungi-y one 
was seen to eat 
the larva, which it 
evidently disliked. 



Mr. Butler teUs me 
that these larvae 
were sapjjlied to 
the young of the 
Great Tit by the 
parent birds. On 
the other hand, 
Eev. G. J. Burch 
tells me that chick- 
ens pecked at the 
]arva3, but rejected 
them, and that the 
hen warned them 
against such food. 



How far evidence 

supports 

Wallace's suggestion. 



How far evidence 

supports 

Poultun's suggestion. 



Couiplete support. It would 
ha\e been very interesting to 
have also experimented with 
the Lizards. 



It would be interesting to also 
try Birds and other smaller 
Lizards. This experiment 
alone opposes suggestion. 



Support ; for the unpleasant 
qualities certainly produced 
eli'ect in some cases. 



Support, from the behaviour of 
the Lizards and from Mr. 
Burch's observation. 



Complete support. The work- 
ing of a counter principle is 
well seen — the uncertainty of 
the birds, their evidently 
half-detecting suspicion, and 
finally the trial wliich soon 
proves the impo.sitiou ; all 
this shows the danger ot reli- 
ance on such a luode of de- 
fence. 



Support, if it be found that the 
appearance is really terrifying 
to enemies, and yet that the 
Lizard ate the larva through 
hunger. 



Strong support from the beha- 
viour of the Lizard, which 
ate the larva reluctantly, 
although disliking it, because 
it was hungry. 



It is probable that they would 
have been eaten, like the 
Croesus, if the Lizards had 
been very hungry. Mr. But- 
ler's birds appeared to relisli 
the larvaa. 



15» 



208 MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 

Table III. — Not Inconspicuous Larvce, which are 



Species 

of 
Larva. 



Pajji/io-nia- 
chaon. 



Method by which 

rendered more or less 

conspicuous. 



Unpleasant 
attribute. 



Arctia caja 



I'hragma- 
tohiafuli- 
ginosa. 



Spilosoma 
meiithas- 
tri. 



Lasiocam- 
pa rubi. 



Green, black, and orange. 
"A striking appearance" I 
( Weinmann). Granting 1 

that this is so, when the 
larva is looked at alone I 
think that its colours har- ' 
nionize well with its umbel- 
liferous food-plants. How- t 
ever, when disturbed the ' 
pinkish-red everted glands 
do render it conspicuous ; 
but this is afier discovery. 



A pair of dorsal pro- 
thoracic glands, 
everted when an 
enemy approaches, 
and causing a most 
penetrating odour 
like " decaying pine- 
apple " (Buckler) ; 
especially noticed in 
fourth stage. 



Eesults of 



J. Jenner Weir. 



A. G. Butler. 



Black, with very long grey [ Jenner Weir thinks 
and brown hairs. Freely ' that " flavour is 
exposed on dead nettle, &c., ] nauseous " and that 
although the colour is not the hairs act as a 
such as to at once attract warning. Neverthe- 
attention. less I am assured that 

the shorter hairs are 
irritating ( W. Cole). 



Smoky brown, with brown 
hairs. Freely exposed on 
dock &c. ; as above, it is 
not truly conspicuous. 



Lasiocam- 
pa quer- 
cus. 



Odenestis 
J oiaioria 



Brown, with lougbrown hairs. 
Freely exposed on all low 
plants ; as above. 



Black and brown, with long 
brown hairs. Freely ex- 
posed on heather &c., as 
above ; more conspicuously 
coloured with black aud 
yellow bands when younger. 



Evidence, as far as it 
goes, against there 
being any unpleasant 
attribute, but only 
tried with Lizards. 



In this case much evi- 
dence for the larvse 
having unpleasant 
taste. Jenner Weir 
thinks hairs are 
warning. 



I believe that there is 
evidence for the hairs 
having irritating 
pi-operties. 



Disregarded by all 
the birds. 



Young and compa- 
ratively hairless ; 
tasted and refused 
by many birds ; 
disregarded by all 
when older aud 
very hairy. 



Devoured by L. viri- 
dis. 



Brown and yellow, and com- 
paratively conspicuous when 
young ; black with white 
marks and brown and grey 
hairs when older; exposed 
but not conspicuous on 
hawthorn. 



I believe that there is 
evidence for the hairs 
having irritating 
properties. 



Disregarded 
the birds. 



by all 



Blue-grey, black, and orange; 
tufts of white hairs; freely 
exposed on grasses. Easily 
seen, but does not attract 
attention. 



The hairs are cer- 
tainly irritating, al- 
though it takes some 
time to affect the 
hands. 



Disregarded by all 
the birds. 



188".] VALUE OF COLOUR AND MARKINGS IN INSECTS. 209 

ifot Nocturnal and which do not conceal themselves. 




210 MR. E. B. POOLTON ON THE PROTKCTIVE [MhF. 1, 

distasteful species are carefully excluded from the diet. Thus 
Butterflies and Moths are freely eaten by Lizards (see Appendices) ; 
but I am sure that tliey are not really enjoyed in the same way as 
when a Housefly or a palatable Caterpillar is offered to them. This 
is doubtless because the imagos of Lepidoptera are dusty, unsatis- 
factory things to eat, with such a small proportion of body in 
which the real nutriment and taste is contained, and so large an 
expanse due to the dry membranous wings with their scaly 
covering. In this respect the Butterflies contrast unfavourably (as 
food) with the Moths, and the latter are certainly preferred (when 
both are palatable in other ways). The same preference is manifested 
by Frogs {Hyla nrborea) with even greater force ; there is a most 
extraordinary difference in the behaviour of such a Frog in the 
presence of a Housefly or of a Butterfly respectively, and in fact the 
latter is often disregarded. Of course birds are in a different position 
as regards such insect-food, for they at aiij' rate very generally 
pick off the unpalatable parts before eating a lepidopterous imago 
(Jenner Weir) ; and with them it is common to witness all the signs 
of an intense desire for these insects, especially Moths. Birds can 
similarly largely remove the unpleasantness due to larval hairs, as 
was seen in the case of 0. antiqua (Table 1.). We should doubt- 
less see evidence ibr the existence of such nice discrimination 
between the relative palatabilities of various insects, in the case of 
all insect-eaters, if our observations were sufficientlv numerous and 
minute ; but it must be quite clear that the preferences cannot 
be always satisfied, when we remember the extent and keenness of 
competition. In this country it is hard to realize the excessive 
abundance of reptile life, chiefly among the Li/ards, which obtains 
even so near to us as the south of Europe, and which almos^t entirely 
depends upon the insect fauna for food. Almost every step along 
an Italian road startles several Lizards on the road-side wall or bank ; 
and it must be perfectly clear that under such circumstances it is 
quite impossible for all to be served with the food which is most 
appreciated. We see rather the very conditions which must render 
the acquisition of an unpleasant taste together with the correlative 
"warning" colours, an exceedingly hazardous mode of protection, 
if assumed by more than a small propoition of the species constituting 
the insect fauna of such a country. For in so great a press of 
competition among the itmumerable insect-eaters, we may feel sure 
that some at least would be sufficiently enterprising to make the 
best of unpleasant food, which has at least the advantage of being 
easily seen and caught. And such a conclusion will, I think, be 
confirmed by a study of the tabulated details. It must be admitted 
that Wallace's suggestion, with its experimental proof, has taken a 
most important place among the princifdes which deal with the 
infinitely complex and ever-changing relations which obtain between 
the most widely separated no less than between the most allied 
members of the organic kingdom. But it is no less true that the 
principle carries with it its own compensating principle, which will 
come into operation precisely as the former advances to the possession 



1887. J VALUE OF COLOUR AND MARKINGS IN INSKCTS. 211 

of undue influence and thus throws out of adjast;nent the preexisting 
condition of comparative equilibrium. 

Another conclusion which is demonstrated very completely by the 
tables is that a Caterpillar may be eaten by one insect-eating Vertebrate 
although refused by another. I believe, however, that the acquisition 
of an unpleasant taste and of conspicuous colours appealed, at any 
rate at first, to a large number, probably all, the vertebrate foes ; 
for if this were not so, if the species became unpalatable and 
conspicuous to (say) half its enemies, and became conspicuous but 
remained palatable to the remainder, it seems only reasonable to 
conclude that immunity from the attacks of one set of foes would 
be counterbalanced, or perhaps more than counterbalance!, by the 
facilities afforded to the other set. On this account and for other 
reasons which will be given below, I thiniv it probable that the 
differences observed between the enemies of insects in this respect 
are of recent date as compared with the acquisition of this mode of 
protection, and have arisen out of the great competition for food ; but 
in most instances the change of habit has not become so far confirmed 
that the previously distasteful food is eaten with avidity and pleasure. 
The first table of highly cons|)icuous larvae (including Crcesus and 
Nematus from tiie second table) can be shortly analyzed to show in 
It the various stages of transition from the most utter disregard to 
the opposite extreme of conduct, indistinguishable from that observed 
when the larvae are known to be relished. The intervening stages 
are furnished by the details given by the different observers, and 
are described in the headings of the vertical columns between those 
numbered I. & V. (see page 2 1 2). 

It must be remembered that these analyses represent a comparison 
between the results of experiments carried out under different systems 
and with the use of an incomplete number of Vertebrates in all 
cases. Hence many of the insects would doubtless have to be 
shifted into other columns after being offered to other Vertebrates, 
or to those actually employed, if it were certain that they were 
thoroughly hungry, Allowing for this, however, the analyses provide 
us with numerous instances of transition through all conditions of 
failure in the protective efficacy of the method we are discussing. 
A.t the same time one can see at a glance the relative behaviour of 
different insect-eaters as far as they have been tested in the case of 
each larva. 

Jenner Weir's suggestion tiiat the hairs of certain larvfe act as a 
warning of other unpleasant qualities can also be tested by the 
examination of the former tables. There are altogether fourteen 
larvae which may be called hairy, out of a total of twenty-seven 
(omitting the two terrifying species). Of these, two (Z. rubi and P. 
Julifftnosa) were eaten, as far as any observations are recorded ; one of 
the former and five others (L. rubi, L. quercus, L. pini, P. auriflua, 
A.caja, and O. potatoria) are either known to possess irritatin* 
hairs or are believed to possess them ; as many as five are gregarious 
(r. 20, V. urticfB, P. bucephala, E. lanestris, C. neustria), and this 
habit, together with the colour, is by far the most important factor 



212 



MR. K. U. POULTON ON THIC PRdTKCTIVK [Mar. 1, 



B. = Birds, F. = Frogs, L.=;Lizarcls, S.=Spider3. 



A. Disregarded by all Vertebrates I V. io, A jillprndiila;, D. ccsru- ^ 
as far as experiiueuts have -^ Icocephala, H. defolinria. | 
been made at present. | D. (/alii, P. hrassiccs 



B. 
L. 



B. Disregarded by some foes ; f 
tasted and rejected by others. \ 



f C neusfria , B., L. 

I E.jacoheecs L. 



C. Disregarded by some foes, 



A. grossidariata ... 
P. huccphala 
0. antiqna 



out eaten, m some cases with-i n , ", . 
rehsh, by others. 



P. miriflua 
G. verbasci 
V. urfice... 
^ N. ribtv/'i . . . 



B.,L.,F 

L. 

B. 

B. 
B., L. 

B. 

B. 
B., L. 



D. 



Disregarded by 
enemy. 



no species 



of 



C. f,epfcnfrionaUs 
H. wavaria 



E. Eaten witli more or less pleasure { L. pini 

by all the species of enemy on \ 

■which experiments have been [ i). euphorbia; 

made. 



II. 






33 



L. 
L. 

F.,S.,L 
L. 



L. 

b! 

L. 

R, S. 



III. 



k-.JH 



id >^ 



W 



L. 
?L. 
L. 
L. 



IV. 






B. 
L. 



V. 






?B. 
?B. 
L. 
B. 

L. 
B. 



B.,L. 



Similarly analyzing the third table, we find the following results : — 





I. 


II. 


III. 


IV. 


V. 


1 p. rnachaon 


L. 

B. 

B.,L. 

B. 

B.,L. 


B. 
L. 


L. 




?L. 
?L. 


A. \ A. caja, S. ■menthasfri (old) 
1^ L. quercus 


B. S. menthasfri (you ng) 


C. 0. 'potatorm 

-p { P. fidic/inosu 


' 1 L. n(bi 





1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 2\'A 

ill producing a conspicuous appearance, although it may be admitted 
that the hairs do render subordinate assistance ; of the two remaining 
larvae, one is brightly coloured (O. untiqiia), although the hairy tufts 
are in this case Aery important factors, while for S. menthastri no 
other suggestion except that of Jenner Weir has yet been made. In 
some of these larvae the effective colours are cluefiy on the hairy 
covering, and the latter practically makes up the whole appearance. 
Turthermore in the last species there is evidence (Jenner Weir) that 
the insect is dishked for some quality other than the presence of 
hairs. It seems probable that the hairs of larvae possess irritating 
qualities to a much greater extent than is commonly supposed ; but 
it is also likely that the hairy covering may be of direct value to the 
organism in other ways, some of which doubtless remain to be dis- 
covered. (Is it not likely that some tactile or other terminal organ 
of the nervous system may be in relation with hairs or bristles ?) 
Of course it is well known that hairs are often exceedingly important 
in defending the insects by the converse method of a protective 
appearance (an extreme instance of this is afforded by the larva of 
Acroriycta leporina, see Trans. Ent. Soc. Lond. 1886, pt. ii. June, 
p. 160). 

Prof. Meldola has kindly looked through the jiroof-sheets of this 
paper, and has suggested to nie that the probalde original significance 
of tlie hairy covering was protection frtnn injury after fiUling from 
the food-plant. The suggestion is strongly supported by the habits 
of the majority of hairy larvae. 

Any further considerations which arise out of the tables given 
above are better deferred until after an examination of similar 
instances in forms other than the larvae of Lepidoptera. 

II. Briglitly coloured or conspicuous Insects {other than LarvcB). 

Under this heading I have only included such species as have 
been actually subjected to experiment. There are a very large num- 
ber of additional species from many groups of insects which doubtless 
belong to this category ; but as they have never been brought to the 
experimental test, they are excluded from consideration on the present 
occasion. An extended list will be found in Wallace's ' Contributions 
to the Theory of Natural Selection ' already referred to (The essay 
on " Mimicry and other Protective Reiemblances among Animals"). 
In most of the instances given by Wallace, we may feel confident 
that the test would prove satisfactory, especially as the author shows 
that in many cases the conspicuous form is mimicked by one or more 
species belonging to totally different groups, which accompany the 
former in its range and which, nearly always, keep in a small minority. 
Such facts render it in the highest degree probable (in fact make it 
nearly certain) that the mimicked species possesses some excep- 
tional advantage in the way of inedibility or otherwise — some 
unpleasant peculiarity which confers upon it a more or less com- 
plete immunity from the attacks of the Vertebrate foes of its class. 
But in the present paper I am rigidly limiting myself to instances 



214 MR. E. B. POULTON ON THK PROTECTIVE [Mar. 1, 

which have been actually tested, and it is much to be regretted 
that experimental investigations have not been further extended 
and recorded in greater detail. The results of the tables of larvae 
given above have been, in a very small proportion of cases, so 
directly contrary to a priori expectation that I do not feel confident 
in bringing forward any instances which have not been tested, although 
I feel sure that the vast majority of them would yield favourable 
results. I cannot, therefore, in this paper accept as satisfactory 
the purely negative evidence that insect-eating Vertebrates have 
been often seen to catch and eat insects of various kinds, but have 
not been seen to catch at the same time and place certain highly 
coloured species which were abundant and slow-flying. At the 
time when Bates and Wallace first made public their most impor- 
tant conclusions as to the meaning of conspicuous coloration and 
the true significance of mimicry, it was quite right that evidence of 
all kinds should be brought forward ; but after the lapse of twenty 
years, we may fairly expect that conclusions which are so important 
in Biology shall have received the most abundant and complete 
experimental proof. And I know that lack of detail in the proofs 
which have been afforded, and the fact that a large part of the evi- 
dence brought forward is still founded on mere surmise (however 
probable may be the result of an actual test), have prejudiced the 
conclusions in the minds of many distinguished biologists, who 
have come to look upon the whole subject with an undeserved 
suspicion. 

I cannot find any record of actual experiments conducted upon 
the well-known and conspicuous Heliconians and Danaids, and 
therefore I do not include them in the following list. There is, 
however, an observation of Meldola's which is of the nature of 
demonstration, and which is so interesting that I quote it in his 
words : — " It appears that tlie nauseous character of these .... 
butterflies is to a certain extent retained after death, as I found 
that in an old collection which had been destroyed by mites, the 
least mutilated specimens were species of Danais and Euplcea, 
genera which are known to be distasteful when living and to serve 
as models for mimicry, see Proc. Ent. Soc. Lond. 1877, p. xii." 
(Meldola's editorial notes to his translation of Weismann's Essays 
above referred to, p. 337). This observation (since confirmed by 
J. Jenner Weir, ' Entomologist,' vol. xv. 1882, p. 160) has the same 
kind of interest as that of Butler upon spiders, drawing attention, 
as it does, to the possession of a peculiar taste or smell which is 
recognized as nauseous by animals as widely separated as the mites 
and spiders are from lizards and birds. And such a consideration 
enforces the conclusion previously arrived at from other evidence, 
that when certain insect-eaters neglect the attributes which are 
respected by others, we see the results of an "acquired taste" 
produced in the first instance by hunger, and not by an obe- 
dience to the dictates of an eccentric preference for what is very 
universally regarded as disagreeable. 

Since the above was written, my friend and pupil Mr. E. A. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 215 

Minchin has called my attention to some experiments made by 
M. (le Niceville upon tlie images of Acrseinse &o. (Butterflies oV 
India, Burmali, and Ceylon, vol. i. part ii. p. 318). " M. de Nice- 
ville has experimented with the carnivorous Mantis on many of the 
Butterflies believed to be offensive to birds, and he has found that 
A. violcB is the only Butterfly which all the species of Mantis he has 
experimented with refuse to eat." 

One other consideration remains to be partially discussed before 
giving the list of experiments. The meaning of conspicuous colora- 
tion may be said to be clear and definite in the case of larvae, being 
only capable of the explanation that it is of value for protective pur- 
poses. But in the perfect forms the other explanation of colour is 
always possible, i. e. its use in courtship, and, as a consequence, its 
growth from small beginnings and its gradual perfection by sexual 
selection. By what criterion, then, is it possible to judge of the 
meaning of bright and cons[)icuous colours in any particular instance? 
In the first place, the brilliant tints due to sexual selection alone 
rarely usurp the whole surface of an insect, and tliere are certain parts 
(exposed in the protective attitude of rest) in which we expect to find 
such brilliant tints replaced by the (usually) sober colours which 
harmonize with the environment. For instance, this is well known 
to be the case with the uppersides of the upper wings in most 
moths, and with the undersides of both wings in nearly all butterflies. 
If, therefore, such exposed parts are cons()icuously coloured, strong 
presumptive evidence will be afforded for the explanation of the 
colour as belonging to the "warning class," Another test is 
iouud in the mode of flight, which may be expected to be such as 
will clearly display the colours no less than when the insect is at 
rest. The question is no doubt complicated by the two kinds of 
coloration running into each other. Thus Wallace has shown 
that the shapes and colours due to sexual selection may run riot in 
localities (certain islands) where enemies are largely" excluded by 
barriers, and in the same way the brilliant colours of dangerous or 
nauseous insects may perhaps be sometimes exj.lained by equal 
immunity, although due to other causes. But a conspicuous appear- 
ance gained in this way will be always liable to be used for an 
entirely different object as " warning coloration." But if such a 
change of use took place, we should expect some change of pattern 
or some new combination of colours, for this reason, that "warning 
colours " have one meaning which is not associated with sexual 
colours, i. e. the production of a conspicuous appearance and the 
more or less complete subordination of everything to this end. 
Nevertheless it is almost certain that the appearance of any species, 
however specialized for other ends, possesses sexual significance, 
and appeals as an adornment to the modified taste of the individuals 
concerned ; and we have a rough criterion of the extent of the 
modification in taste, when we compare such appearances with those 
which possess a sexual value alone, which are concealed except 
during flight and which are especially displayed during courtship. 
Prof. Aleldola has recently drawn my attention to an observation of 



216 Mil. E. B. POULTON ON THE PROTECTIVE [Mar. I, 

Darwin's which enables us to point towards the purely sexual value 
of one factor in the appearance of certain butterflies ; and by the 
same kind of observation it is likely that increasing stability will 
be given to tlie whole theory of sexual selection. Darwin noticed 
that the magnificent surface-colours which exist on many tropical 
butterflies, and which change with the position from which the 
insect is observed {being probably due to diffraction), become 
most brilliant when they are seen from the front, and at an angle 
which corresponds with that at which they would be seen by the 
female as the male butterfly approaches her. 

As a further test of the " warning " value of certain colours, we 
can use as a comparison the colours and patterns of certain larvae 
which are known to be "warning" only, or which at any rate cannot 
be sexual. After giving the list and the experimental details, I 
shall attempt to show that there are certain combinations of colour, 
and sometimes even certain patterns, which are generally distinctive 
of "warning" as apart from sexual coloration. And such dif- 
ferences of type are to be expected ; for the two systems of coloration 
appeal to a different class of animals and appeal to a different 
sense. The " warning " colours of insects appeal to a Vertebrate' s 
sense of what is conspicuous ; the sexual colours appeal to an 
Invertebrate's sense of what is beautiful. And it is one of the 
most remarkable facts in the study of comparative psychology 
that our own sense of what is beautiful entirely coincides with that 
of an insect, so that the difference in the two types of coloration 
depends entirely upon the sense alluded to above, and has probably 
no reference to the class of animals in which the sense exists. 
For, if we had no knowledge of tiie use and meaning of the colours 
of insects, we should all agree in maintaining that certain colours 
and patterns (whicli we now trace to the action of sexual selection) 
comprise all that is beautiful in the appearance of this class of 
animals, and we should draw a sharp line between these and 
other combinations of colour which we now recognize as possessing 
an entirely different significance. 

In the following instances it is unnecessary to enter upon any 
preliminary account, as the table contains all the details which 
I can find recorded. Of course this list must contain any cases 
(if tested) in which a conspicuous appearance is believed to be 
due to " mimetic " resemblance to another species protected by 
possessing some unpleasant attribute. I much regret that hardly 
any of these most interesting cases have been used for experiment 
(see Table IV., pp. '218-223). 

Looking at this fourth table, we find that the theory of warning 
colours is again confirmed in the most marked manner. Unfortu- 
nately the extremely interesting " mimetic " cases still lack the 
necessary experimental demonstration; although the intimidating 
larvae of C. elpenor and C. vinula are really mimetic of vertebrate 
appearances, and the former has been shown to be attended by a 
considerable amount of success. The only "mimetic" form in 
this last list is Sesia fuciformis, and in this case the mimicry is 



1887.] VALUE OF COLOUR AND MAKKINGS IN INSECTS. 217 

exceedingly imperfect, while very perfect protection is gained in 
other ways. In fact I have suggested above that the species is 
not now "mimetic" at all, but retains two points of an ancestral 
condition in which it actually derived shelter from the reputation 
possessed by many Hymenoptera, these being (1) a structural 
point — its transparent wings, and (2) a habit — its diurnal flight. 
If this view be correct the species should be moved into the table 
given below of the results of experiments upon insects protected 
by evading their enemies, and there it would add to the instances 
which support Wallace's converse suggestion, that such insects 
will be found to be eaten just as the class we are discussing 
are generally refused. I have placed the species in the fourth 
table in deference to Wallace's opinion on the kindred species 
S. bombyliformis, which it closely resembles ; but I believe that 
the place of both species should be below. I made several 
attempts to obtain the imagos of Sphecia apiformis, in order to offer 
these to my Lizards ; for this species certainly does suggest a 
stinging Hymenopterous insect in the most remarkable way ; but 
the attempt was unsuccessful. S. bembeciformis is equally well 
suited for experiment, but even more difficult to obtain ; I hope, 
however, to be able to experiment with one or both species in the 
course of next summer. 

It is noteworthy that (excluding S. fuciformis) there is probably 
no instance in this list which proves a difficulty in the way of 
Wallace's suggestion at all equal to that raised by the larva of 
D. euphorbice. For the pupa of A. grossulariata was not swallowed, 
but only well bitten by a very hungry lizard, and although the 
imagos of S. lubricipeda and P. uiiriflua were eaten (also by very 
hungry lizards in the former case), it cannot be asserted that they 
were eaten with relish ; and, further, the experiment should be 
repeated with Birds and other species of Lizards. In all other cases 
the insects were refused by at least one of the animals to which they 
were offered. The comparison between the treatment received by 
Bees (described below) and Wasps, together with a comparison 
between their appearances, is strongly in favour of Wallace's sug- 
gestion. It now remains to analyze the list in the same manner as 
was adopted before, showing the transition of conduct observed (see 
Table, page 224). 

Comparing the three analyses of all experiments ever conducted 
upon presumably nauseous or dangerous insects, omitting S. fuci- 
formis and including C. elpenor and C. vinula, we find that out of a 
total of 44 cases which have been tested 13 were entirely disregarded ; 
but this number includes P. machaon, which I believe ought to be 
placed below under forms chiefly defended by protective resemblances, 
and also S. menthastri (larvte, which were tasted in the youngest 
stages). Furthermore four species (including the two just men- 
tioned) belong to Table III., containing larvae which can hardly 
be called conspicuous. Hence, omitting this table, the number is 
reduced to 9 cases out of a total of 37 ; and of the 9, not a single 
species has been offered to more than one out of the three groups of 



218 



MR. E. B. POULTON ON THE PROTKCTIVE [I\Jar. 1, 



Table IV. — Bright-coloured or Conspicuous 



Species 

and 

Stage. 


Method by which 

rendered more or less 

conspicuous. 


Unpleasant attribute, 
I'eal or suggested, 
in mimetic forms. 


Results of 


J. Jenner Weir. 


A. G-. Butler. 


Pupa of 
Abraxas 
grossula- 
riata. 


Black, with yellow bands ; 
contained " in a very slight 
and perfectly transparent 
cocoon " (Newman). 


? Taste or smell. Proof 
lies in what follows. 








, 


Iiuago of 
Aiifhrocera 

Jilipendu- 

Ice. 


Blackish green and red. 
Rests rery conspicuously on 
flowers &c. ; fl.ight not ra- 
pid, and by day. So abun- 
dant locally as to be almost 
gregarious. The bright 
colours are well seen at 
rest and in flight ; very 
sluggish and easily caught ; 
often feigns death when 
captured. 


? Taste or smell. Proof 
lies in what follows. 


" Unwillingly eat- 
en:" Jenner Weir 
thinks because the 
birds in confine- 
ment get fewer 
insects than when 
wild, and so wiU 
eat forms which 
they would reject 
if well supplied 
with this food. 


Always refused by 
L. viridis after 
having been 
seized. 


Imago of 

Sesia fu- 
oiformis. 


Wings transparent, with 
brown margin ; body vari- 
ous shades of brown. Flight 
by day veri/ rapid, and in- 
sect seldom seen at rest. 
Its claim here depends on 
its somewhat Hymenopter- 
ous appearance : the closely 
allied S.hombyliformis "re- 
sembles the male of the 
Humble-Bee (Bonibus hor- 
torum)" (Wallace). 


The suggestion of a 
sting, according to 
some authorities. 








j 

1 


Imago of 
Porthesia 
auriflua. 


Almost all the surface, seen 
at rest, is white ; there are 
also a few small black 
patches on the fore wings, 
and the body ends posteri- 
orly in a conspicuous yel- 
low tuft. Flight weak in 
the evening, when the Moth 
is very conspicuous, appear- 
ing white ; at rest it is also 
extremely conspicuous. 


There may be an un- 
pleasant taste or 
smell, but there is 
at present no expe- 
rimental proof of 
this. 


A single imago eat- 
en by the Lizards. 




1 

i 
1 
1 
1 
1 



1887.] 



VALUE OF COLOUR A^D MARKINGS IN INSECTS. 



219 



Insects {other than Larvce). 



Experimenta. 



A. Weisiniinn. E. B. Poulton. 



One was placed in 
cage of Tery hun- 
grj L. mtiralis, and 
was crushed and 
most of the fluid 
contents extracted 
and probably eat- 
en ; howeyer, pala- 
table pupa; were 
swallowed as a 
whole. This pupa 
was untouched at 
first. 



One specimen eaten 
by L. muralis; no 
apparent hesita- 
tion or caution in 
capture. 



How far evidence supports 

the theory of 

"warning" and "mimetic" 

colours (Bates and Wallace). 



Strong support, for if palata- 
ble it would have been 
treated very differently. 



Strong support, from the re- 
luctance of Birds and rejec- 
tion by Lizards. 



At first seems to be a difficulty ; 
but I have always thought 
that the suggested resemblance 
is very imperfect, and perhaps 
a remnant of a former more 
perfect mimicry, reliance be- 
ing now placed on powerful 
flight and concealment during 
rest. 



Bearing of evidence on 

Poulton's suggestion, 

as before. 



Strong support. 



Strong support in Jenner Weir's 
explanation of the Birds' be- 
haviour, and some support in 
tlie Lizard's attempts. 



Insufficient evidence ; but op- 
posed to the theory as far as 
it goes. 



Ko evidence, the insect being 
palatable and in this instance 
not terrifying. 



Ko evidence, unless it should 
be proved to be a nauseous 
insect. 



220 



MR. K. B. POULTON ON THK PROTECTIVE [Mar. 1, 



Table IV. 



Species 

and 
Stage. 


Metliod by which 

rendered more or less 

conspicuous. 


Unpleasant attribute, 
real or suggested, 
in mimetic forms. 


Results of 


J. Jenner Weir. 


A. G". Butler. 


Imago of 

Spilofoma 

menthastri. 


Wings creamy while, with 
black dots ; body yellow, 
with black dot.=. Very con- 
spicuous at rest or in flight 
(slow ; evening) ; often 
feigns death when cap- 
tured. 


? Taste or smell. Proof 
lies in what follows. 


Eaten reluctantly 
by Eobin and 
Eeed-Bunting, the 

latter after reject- 
ing it at first. Tas- 
ted and rejected 
by Yellow-Ham- 
mer ; refused by 
BuUiinch and 
Chaffinch. 







Imago of 

Spilosoma 
lubricipe- 
da. 


Buff, with black spots. Very 
conspicuous at rest or in 
flight (slow ; evening) ; often 
feigns death when cap- 
tured. 


?Taste or siuell ; but 
there is at present 
no experimental 
proof of this. 




' 




1 


Imago of 
Euchelia 
jacobcECB. 


Fore wings very dark brown, 
almost black, wil;h red 
spots and stripes ; hind 
wings red with black mar- 
gin ; body black. IMost 
conspicuous at rest, or espe- 
cially in its weak flight by 
day. 


? Taste or smell, or 
very probably from 
being indigestible. 


Disregarded for 
some time, but 
eventually reluc- 
tantly swallowed 
after the removal 
of the wings. 
Same suggestion 
as in case of A. 
filipendulce. 






Imago of 
Abra.vas 
grossula- 
riata. 


White, black, and yellow. 
Very conspicuous at rest 
or in its slow flight by day 
and in evening. Often 
feigns death when cap- 
tured. 


■/Taste or smell. Proof 
lies in what follows. 


Refused by Lizards, 
although seized on 
one occasion. 


•' Greedily devour- " 
ed " by Frogs. (I 
think that this 
umst be a mis- 
take ) 


Imagos of 
Malaco- 
derms of 
the genus 
Telepho- 
rits, sp. ? 
(Coleopte- 
ra). 


Black and red. Very con- 
spicuous at rest or in 
flight (diurnal) ; easily 
caught. Elytra soft and 
no protection. Common 
English species called "sol- 
diers and sailors," 

i 


? Taste or smell Proof Disrefrnrrlpfl bv nil 




lies in what follows. 

1 


the birds. (Quoted 
by Wallace.) 


1 



1887.] 



VALXiE OF COLOUR AND MARKINGS IN INSKCTS. 



221 



{continued). 



Experiments. 




How far evidence supports 

the theory of 

"warning" and "mimetic" 

colours (Bates and Wallace). 


Bearing of evidence on 

Poultou's suggestion, 

as before. 


A. Weismann. E. B. Poulton. 






Strong support. Also confirmed 
by Stainton, who offered it to 
Turkeys, with a large number 
ofproteetively-colouredMoths, 
all of which were eaten, while 
the S. mcnthadri was always 
rejected after being examined. 


i 
Strong support, especially in 
the reluctant way in which it 
was eaten by Robin &o. Set 1 
Jenner Weir's explanation i): 
case of A.filipendulcB. 

1 








Eaten at once by 
hungry L. muralis. 


! 
So far as it goes, against sug- > No evidence, unless it shoulc 




gestion; but not tried with 
sufficient number of enemies, 
nor with ]jlenty of other 
food. 


be proved to be nauseous ; ' 
then this experiment woult" 
strongly support suggestion. 




Always refused by 
L. viridis. 


Two specimens eat- 
en successively by 
the same Frog, 
and therefore it 
would seem not to 
be nauseous to 
this species. Next 
day the Moths 
were foimd in the 
case, having been 
rejected presum- 
ably because of 
iiidigestibility. 


Strong support. 


• 
Strong support in Jenner Weir's 
explanation as above. In the 
case of the Frog also hunger 
very likely caused the iusecl 
to be taken, although not re- 
tained. 




Eefused by L. mu- 
ralis after biting, 
although very 
hungry. 


Support from the behaviour 
of Lizards ; the other evi- 
dence must, I think, be mis- 
taken. 


Support in that the Lizard: 
were induced to bit.^ it st 
verely. 








Strong 8\ipport. 


No evidence. 


1 




Pr 


DC. ZOOL. SOC. — 


1887, No. XVI. 


16 



222 



MR. K. B. POUITON ON THE I'ROTECTIVK [Mar. I, 



Table IV. 



Species 
and 

Stage. 


Method by wliich 

rendered more or less 

conspicuous. 


Unpleasant attril)ute, 
real or suggested, 
in mimetic forms. 


Results of 


J. Jenner Weir. 


A. G. Butler. 


Imago of 

Chrysome- 

la ■pojmli 

(Coleop- 

tera). 


Chief colour red, due to ely- 
tra : the other parts seen 
are a metallic lustrous blue- 
black. 


"Strongly smelhng" 
{Weisniann). 








1 


Coccinella 
sejJtem- 
punctata 
(Coleop- 
tera). 


Exposed surface red with Tlipv Imve n, verv nn- 


1 : 


black spots. Very conspi- 
cuous and easily caught ; 
flight diurnal. 


pleasant smell. Tliey 
" can emit fluids of 
a Tery disagreeable 
nature" {Wallace). 




1 


Coccinella Exnoserl surface rerl wUli two i Tliev Imve n tpit iin- 






hipvnc- 
taia (Co- 
leoptera). 


black spots. Very conspi- 
cuous and easily caught; 
flight diurnal. 


pleasant smell. They 
"can emit fluids of 
a very disagreeable 
nature" {Wallace). 


', 




Vespa vul- 
garis. 


Queens and workers made use ; Stino-. 






of: colours black and yellow. 
Very conspicuous ; power- 
ful flight, but of use, for 
obtaining food and catching 
prey rather than for esca- 
ping enemies. 








Bomhus 
terrestris, 
B. lapi- 
darius, 
&c. 


Other common species also 
made use of. The msects 
are generally conspicuous, 
with bands of light colour 
(often yellow) on a darker 
ground (often black). Their 
size also attracts attention, 
while their flight is clumsy 
and lieavy, and they do not 
rise quickly when at rest. 


Stings. 


Bombus ? sp., killed, 
but not swallowed, 
by L. viridis. 


Bombus terres is 
eaten by L. viridis, ' 
after cautious dis- 
ablement, as in 
the case of Bees. 
Only eaten when 
hungry. 


Nomada 
marska- 
mella. 


Very conspicuous, with black 
and yellow bands. 


Sting. 











1887.] V.VLUE OF COLOUR .\ND MARKINGS IN INSECTS. 223 



(coiiiinued). 



Experiments. 


How far evidence supports 

the theory of 
" warning " and " mimetic" 
colours (Bates and Wallace). 


I 

Bearing of evidence on 

Poulton's suggestion, 
as before. 


A. Weismann. 


E. B. Poulton. 


Always rejected by 
L. viridis. 




Strong support. 


No evidence. 






Refused by Frogs 
without tasting ; 
also by Lizards. 

Miss Cuudell tells 
me that Hj/la arho- 
rea will eat Lady- 
birds in the win- 
ter when food is 
scarce. 


Strong support. 


Strong support from Miss Cun- 
dell's observation. 






Refused bv Froi^s ' Strnnop sjinnnrt. Wnllnf^A nlso 


Strong support from Miss Cun- 
dell's observation. 




without tasting. 
See Miss Cundell's 
observation quoted 
above. 


says of the Coccinellidte, to 
wliich family this and the last 
species belong: — "Certainly 
rejected by some birds;" but 
no details are given. 




Three Frogs in suc- 
cession caught one 
queen Wasp and 
then released it 
very quickly, and 
after that it was un- 
touched. Lizards 
would not touch 
it, but watched it 
narrowly. 


Strong support. Romanes also 
told me he had seen a Spider 
capture a Wasp, and its great 
caution in the process was a 
further support, as also is the 
fact that Spiders generally re- 
lease Wa.sps from their webs. 


The Frogs went as far as or 
farther than could have been 
expected. 






Bombiis lapidarhiK 
eaten by L. viridis, 
but I have not 
witnessed the me- 
thod. Refused on 
many occasions. 


Support, upon the whole ; they 
would certainly not have been 
touched by the Lizards if 
there had been abundance of 
other food. 


Strong support for an analo- 
gous suggestion for insects 
protected by stings instead of 
)y a nauseous taste. 






Untouched by any 
of the Lizards. 


Strong support. 


This seems to indicate that the 
fear of a sting is very strong; 
for when the insects are too 
active to be killed without 
stinging (also the case with 
Wasps), they are not touched 
by the Lizards. 





16^ 



224 



MR. F. B. rOULTON ON THE PROTECTIVE [Mar 1, 



ei 
be 



( Chrysoviela popuU j L. 

. 1 Malncoderms of the genus Tele- 



pJiorus 
Nomada marshamella. 



B. Vespa vulgaris 
( A. fiUpendulce 



B. 
L. 

L.,F. 

L. 



A. grossulariata L. 



0. 



5. menthastri B. 

E. jacoh(B(B L. 

Boinhns terresfris, B. lapidarius) 
&c •. L. 

Coccinella hipunctata 1 F. 

^ Coccinella septem-punctata I F-.L- 



D. 



E. ^ 



'' Pupa of A. grossulariata 

S. luhricipeda 

S.fuciformis 

P. auriflua 



IT. III. 



% 



a 

(1) 
.^ 
en 

H 



F. 

L. 
B. 



IV. 



i f^-9 I 

3 O I 

— 3 i 

s| i 



3 



33 



.,.( 



B. 

B. 
B. 

L. 
F. 
F. 



L. : 
or II. 

L.? 

orlY. 
or V. , 



V. 



a 



F.?(I 
think 

a mis- 
take). 

F. 

(at the 
time). 



L.? 
or W. ! 
or III. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 225 

insect-eaters. Again, only one species out of the 9, viz. Nomada 
marshamella, always remained untouched by very hungry animals 
when other food was withheld ; and we can only surmise as to what 
would have been the results if the other 8 had been similarly tested. 
It is, however, quite certain that many of them would have failed. 
This is, indeed, proved by the following figures: — Out of the 37 
cases 15 were exposed to this rigid test, i. e. the species in the columns 
under my name in Tables I., II., and IV. (excluding S.fuciformis and 
C. neusiria), and of the 15 only three remained untasted, and of 
these two have been shown by Miss Cundell to be eaten under cer- 
tain circumstances. 

Looking at all these figures, and especially the last, we can well 
understand the following objection being urged against Wallace's sug- 
gestion. It may be said, the tables, indeed, show that Wallace was 
right in predicting that an unpleasant attribute would be found to be 
associated with a conspicuous appearance. That has certainly been 
proved by the results of a vast majority of the experiments ; but of 
what value is this association when insects are seized, tasted, and 
rejected in spite of the warning colours which, ex hypothesi, are 
assumed to prevent this very contingency ? In the first place, an 
answer to the objection is found in the very fact that the insects 
were tasted and rejected to a much greater extent when the verte- 
brates were thoroughly hungry, for we see that when other food is 
present the conspicuous insects are, as a rule, untasted. We shall 
presently see that an inconspicuous but nauseous insect is approached 
by Lizards in a very different way from one which is cons])icuous 
and nauseous. There was, in fact, strong evidence in the details of 
some of my experiments, that the vertebrate enemies were well 
aware that the insects were distasteful, and yet, when exceedingly 
hungry, did their best, in some cases successfully, to eat them. 
This was especially seen in the behaviour of the Lizards towards 
the larvae of O. potatoria, P. bucephala, and Crcesus septentrionalis 
in rny experiments, and towards the larvse of C. neustria, A. gros- 
sulariafa, and P. aurijlua in Jenner Weir's experiments ; for in 
all these cases the Lizards made repeated attempts to eat the larvae, 
again and again rejecting their prey with every sign of disgust 
(rubbing the month against the cage) ; and yet in the end the 
larvae were reluctantly eaten. I believe, however, that the suspicion 
with which conspicuous insects are approached results from the 
strongly impressed experiences of early youth and not from a habit 
which has become hereditary. In many cases, however, the warning 
experience may have been gained without tasting the insects ; for 
we have seen that the latter are often protected by smell, which 
can be perceived from a distance. Excluding these instances, 
however, the experience of conspicuous nauseous forms must have 
been gained by actual trial of a large number. I hope to be able 
to show that it is not necessary for the young insect-eating 
Vertebrate to actually make trial of every species of unpleasant- 
tasting insect in its locality, in order to be equipped with an effi- 
cient stock of experiences with which to conduct its later life. 



226 MR. E. B. I'OULTON ON THE PROTECTIVK [Mar. 1, 

Such an education would be somewhat dearly bought; it would be 
unpleasant to the insect-eater and destructive to the insect. 
But if, as I shall endeavour to show, there is a superficial resem- 
blance between the colours employed by very different insects, 
and frequently even a similarity of pattern, we see that a com- 
paratively few unpleasant experiences would be sufficient to create 
a prejudice against any insect with colours and patterns at all 
resembling the nauseous forms which have already produced so 
indelible an impression upon the memory. And thus it is most 
probable that the conspicuous appearance which astonishes one 
sense becomes associated in the mind of the Vertebrate insect-eater 
with the well-remembered effect of other qualities upon other senses. 
Different Vertebrates vary much in their rates of education. Thus 
my Frogs were much more stupid in this respect than the Lizards ; 
but then the imperfect memory or limited power of generalization in 
Frogs is less fatal to insects than it would be in the case of the other 
Vertebrates ; for I do not think that the larvse were ever injured in 
the least after having been tasted by these animals. It is therefore 
probable that the gradual development of warning colours by natural 
selection was due to the fatalities which followed the experimental 
tasting of other Vertebrate enemies (especially Birds and Lizards), 
which inflict incidental injuries during the process of tasting. But 
the warning appearance having been acquired by such means, the 
Frogs have certainly taken the opportunity (thus offered to them 
read3'-made and without having themselves contributed towards its 
existence) to acquire a somewhat limited education. This was seen in 
the case of the queen wasp (see Appendix IL), which on being placed 
in the case was tasted by three Frogs out of twelve, but afterwards 
■was untouched for nr.any hours (as far as I was able to observe). A 
proof of the limited extent of the education is given by Butler, who 
speaks as if his Frogs repeatedly tried to eat the two species of 
nauseous larvae (A. grossulariata and H. loavaria), seeming only to 
become suspicious when thej' had actually made a trial of the insects 
on each separate occasion. It also seemed to me that my Frogs 
generally, if not always, ate bees from want of memory or deficient 
discriniiiiation ; for in nearly all cases they were finally rejected. 
But the experience did not seem to make any difference to the readi- 
ness with which the next bee would be seized and again rejected. 
On the other hand I did not see a ladybird tasted on any occasion. 
The slight power of discrimination possessed by Frogs was also 
shown by the fact that they frequently jumped at and seized the 
dark-coloured ends of the forceps with which I used to introduce 
insects into their case. 

For the tolerably complete demonstration of the principle which I 
believe has been at work, a far larger number of observations are 
necessary, while complete confirmation requires experimental evidence 
with young Vertebrates which have been reared in confinement, so that 
the whole ( f their education is under observation. As conducing 
towards tliis end, I publish the suggestion with its foundation on the 
resemblances indicated by the tables given below, which have this 



1887.] VALUE OF COLOUR ANO MAKKINGS IN INSECTS. 227 

advantage, tViat they only include insects which have been sub- 
jected to actual experiment. Although the tables comprise so few 
instances, I think that the resemblances of colour and pattern are 
most remarkable, and hard to explain under any other theory. My 
suggestion does at any rate point out a very obvious use for the 
resemblances. The advantages which every conspicuous and nauseous 
or dangerous species would gain by setting as simple a lesson as 
possible to the foes of its class, would be so great that there is no 
difficulty whatever in the supposition that every stage towards con- 
vergence in colours and in patterns would have been beneficial, 
and, as such, would have come under the influence of natural 
selection. It is to be noted that advantage would accrue in the 
greater thoroughness of the education, no less than by shortening 
the process ; for a few colours, with a few simple patterns scattered 
over a number of species, would be remembered more easily than a 
larger number with a separate pattern in nearly every species. 

I am aware that this suggestion is but an extension to the whole 
group of conspicuous insects of the explanation offered by Fritz 
Mviller to a fact which seemed for a long time an inexplicable 
difficulty, the undoubted fact that conspicuous butterflies presumably 
protected in the most complete manner by nauseous attributes, 
nevertheless mimic each other in the most unmistakable way. Bates, 
the original discoverer of " mimicry " in the animal kingdom, pointed 
out these apparently mysterious resemblances in the paper in which 
"mimicry" was itself explained and illustrated. Wallace looked 
upon these obscure similarities between protected forms as due to 
some unknown cause connected with locality. 

It remained for Dr. Fritz Miiller to explain the difficulties in a 
paper entitled •' Ituna and T/ii/ndia; a remarkable case of Mimicry 
in Butterflies" (' Kosmos,' May 1879, p. 100). Arguing from the 
instance of these two genera, which both belong to protected groups 
and wliich resemble each other, Dr. Miiller suggested that under 
these circumstances an advantage would be gained by each of them, 
because the number of species which must be sacrificed to the 
inexperience of young birds and other enemies would be made up 
by both of them instead of by each independently. This paper was 
translated by Prof. Meldola, and appeared in the * Proc. Ent. Soc. 
Lond.' (1879, p. xx). In a subsequent paper by Dr. Miiller 
(' Kosmos,' V. Jahrgang, 1881), the same subject is considered in 
greater detail, and the results are accepted and expounded by Wallace 
in ' Nature ' (vol. xxvi. p. 86). The mathematical aspect of the 
subject was, however, inaccurately stated in this last paper, the correct 
statement being supplied by Mr. Blakiston and Mr. Alexander of 
Tokio, Japan ; the correction being published in letters by Mr. 
Wallace and Prof. Meldola to 'Nature' (vol. xxvii. p. 481). 
Subsequently a letter appeared in ' Nature' (vol. xxix. p. 405) from 
Mr. Blakiston and Mr. Alexander, giving the complete mathematical 
statement of the advantages gained by each of the protected species. 
The law is given in these words, " Let there be two species of insects 
equally distasteful to young birds, and let it be supposed that the 



228 MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 

birds would destroy the same number of individuals of each, before 
they were educated to avoid them. Then if these insects are 
thoroughly mixed, and become undistinguishable to the birds, a 
proportionate advantage accrues to each over its former state of 
existence. These proportionate advantages are inversely in the 
duplicate ratio of their respective original numbers, compounded with 
the ratio of the respective percentages that would have survived 
without the mimicry." 

It had been previously argued that in the case of two protected 
species which had thus come to resemble each other, the proportionate 
advantage was cliiefly on the side of the one which was smaller in 
numbers, and that when the numerical difference was great the 
advantage to the other could be neglected. The amended law wliich 
is quoted above shows, however, that the proportionate advantage is 
always the same, and this is also enforced in another part of the same 
letter: — " It must be remembered, however, that B does no harm to 
A by mimicking it ; on the contrary the act of mimicry is of advantage 
to A over its former state of existence as well as to B ; but A being 
more numerous the advantage is less. Siill, after the assimilation, 
neither has an advantage over the other. Proportionally they suffer 
from tbe ravages of birds equally ; the percentage of losses is the 
same ; they are on equal terms. No matter how long they continue 
the association, neitber gains or loses on the other ; though through 
one being more numerous it loses more individuals, yet equally in 
proportion with the other. So that if one is twice as numerous as 
the other at the time of assimilation, it must always — other condi- 
tions being equal — remain twice as numerous." 

Dr. INIiiller's interpretation was at first criticized in man\' quarters, 
the chief objection brought forward being the belief that birds do not 
learn the meaning of the conspicuous colours by experience, but that 
they avoid such insects by instinct, the ancestral experience having 
become hereditary. There is, however, no direct evidence for this 
view, and I think the account of J. Jenner Weir's observations upon 
Lizards, and my own upon Lizards and Frogs (given in the two 
Appendices to this paper), will go far to furnish an experimental 
refutation of such a theory, so far as these animals are concerned. 
In addition to this, I am assured by a very keen observer. Rev. G. J. 
Burch, that recently hatched chickens certainly do peck at insects 
which they afterwards learn to avoid without trial, and he believes 
that the hen assists in their education by indicating that certain 
insects are not fit for food. His observations were chiefly made upon 
a common phytophagous Hymenopterous larva which is found upon 
gooseberry (doubtless Netnatus ribesii). Another observation made 
by Mr. Burch bears upon the same question. He offered his chickens 
a quantity of chickweed, knowing that this plant was often given as 
food to Linnets. The chickens ate the plant readily enough, but they 
were all extremely unwell in consequence, and vomited freely. After 
this Mr. Burch again offered them chickweed, but they had profited 
by the experience and would not touch it. 

The chief attack upon Dr. Miiller's suggestion was made by Mr.W. 



18S7.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 229 

L. Distant (' Nature,' vol. xxvi. p. 105 ; and ' Rhopalocera Malayana,' 
pt. ii. p. 33) ; but all his objections were very completely answered 
by Prof. Meldola (' Ann. & Mag. Nat. Hist.' Dec, 1882), who, in his 
concludino; sentences, largely anticipates that further extension of 
Fritz MiiUer's theory which is here brought forward, the suggestion 
that all the conspicuous and dangerous or distasteful species in any 
country will be found to share between them a few strongly contrasted 
colours, arranged in few and simple patterns again and again repeated. 
He says : — " I am persuaded that the extension of the theory of 
mimicry proposed by Fritz Miiller marks a great advance in our 
views on this subject, which is so interesting as having been the first 
to which the Darwinian Theory of Evolution was applied with such 
success by Mr. Bates. Not only are we now in possession of a 
consistent theory which enables us to dispense with mysterious and 
'unknown local causes,' but other groups of facts hitherto incompre- 
hensible are capable of explanation. Thus the prevalence of one type 
of marking and colouring tiiroughout immense numbers of species in 
protected groups, such as the tawny species of Danais, the barred 
Ileliconias, the blue-black Euploeas, and the fulvous A^rceas, is 
perfectly intelligible in the light of the new hypothesis. While the 
unknown factors of species-transformation have in these cases caused 
divergence in certain characters, other characters, viz. superficial 
colouring and marking, have been approximated or prevented from 
diverging by the action of natural selection, every facility having been 
afforded for the action of this agency by virtue of the near blood- 
relationship of the species concerned. When discussing the origin of 
mimicry, Mr. Darwin long ago suggested that it might have 
commenced at a time when the species were nearly related in marking 
and colouring." The suggestion here brought forward and depending 
upon the results which are tabulated below, is a further extension of 
the same principles, so that certain resemblances between insects 
belonging to very different groups are accounted for on the 
supposition that natural selection has not only prevented divergence 
in nearly related forms which were originally similar, but has in 
other cases actually determined the convergence of widely separated 
forms which were originally unlike. This latter explanation of the 
resemblances was intended by Fritz Miiller in his paper on " Ituna 
and Thyridia,''^ for he looked upon these genera as widely separated, 
and their similarity as due to convergence. There appears, however, 
to be some dis|)ute as to their true affinities. It is obvious that under 
Prof. Meldola's suggestion we shall expect to find a far greater 
similarity between the species of a large group of closely allied 
nauseous insects in any country than between those of other large 
groups protected in other ways ; while, on the other hand, there is 
no necessity for the expectation of equal uniformity among the 
isolated nauseous species or even among those belonging to small 
nauseous groups. We should rather expect the constant appearance 
of a few simple but very different patterns, made up of a few strongly 
contrasted colours ; and this is precisely the arrangement which is 
proved to obtain by the tabulation of the appearances of all such 



230 Ml». K. B. POULTON ON THE PROTECTIVE [Mar. 1, 

species known to be nauseous or dangerous. Tliere must cer- 
tainly be a tendency towards a further general convergence, but 
the existing condition of convergence round a few well-marked types 
of pattern and colouring must be highly beneficial, and there was in 
this case no initial uniformity due to close affinity, upon which to 
base a general and uniform system. It was in fact « priori far more 
liki-ly that the convergence of remote species should have been round 
a few successful types, while the prevention of divergence among 
closely related species must ipso facto have tended to produce 
concentration round a single tvpe. It will be shown below that 
Fritz Miiller's principle is probably attended by others, which also 
assist in producing convergence, at any rate in some cases. 

Another lesult of the different origin of the two classes of re- 
semblance alluded to above is that the uniform warning colours of a 
large group of closely related species are less conspicuous, and in 
themselves possess less of "warning"' characteristics, than those of 
the smaller groups into which the isolated nauseous species tend to 
converge ; for the former depend largely upon some ornamental type 
of colour and marking, due to sexual selection, and prevalent before 
the time when the nauseous attributes arose. Such a type has no 
doubt been modified in the direction of greater conspicuousness on 
the uppersides of the wings, while bright colours have appeared on 
their undersides, and the mode of flight has been changed into one 
wliich gives the colours their maximum effect ; but still, in spite of 
these changes, the whole appearance of such large groups presents us 
with the ancestral sexually selected colours and patterns, which are 
of great beauty, and are no doubt still of great significance as 
secondary sexual characters. The success of such a stereotyped 
ornamental appearance for warning purposes has depended upon the 
modifications alluded to above, but principally upon the very fact of 
its prevalence and uniformity. On the otlier hand the smaller con- 
vergent groups of nauseous insects often present us with ideally 
perfect types of warning patterns and colours — simple, crude, strongly 
contrasted — everything subordinated to the paramount necessity of 
becoming conspicuous. For the nauseous attributes arising inde- 
pendently among the scattered species of many genera, or in all the 
species of small genera, instead of being chiefly concentrated among 
the m.embers of some one or two dominant groups, it must have 
become impossible to rely upon the slightly altered ornamental 
appearance existing at tiie time when the attributes arose ; but it was 
necessary to appeal strongly to the memory of enemies by the acqui- 
sition of some special form of pattern and colour, in which everything 
is subordinated to the '• warning" characteristics. In the one class 
the pre-existing ornamental appearance was sufficiently well known 
to serve as a warning; in the other class it was not sufficiently well 
known. 

It is quite clear that the tuo classes of resemblance which have 
been just considered must be carefully distinguished from true 
mimicry, in which the mimicking species is without any unpleasant 
attribute, but shelters itself under the reputation of the (nearly 



18?/.] VALUE OF COLOUK AND MARKINGS IN INSECTS. 231 

always) more abundant species which it resembles. In the former 
classes of resemblance we have groups of two or more conspicuous 
forms all possessing unpleasant attributes, which become convergent 
in external appearance, or which maintain an initial uniformity, 
and in either case are mutually benefited by the process. In tlie 
latter class the resemblance would be a source of danger to the 
mimicked species if the edibility of the mimicking species were 
discovered ; and the experiments detailed in the present paper show 
how likely it is that such qualities would be discovered if tlie latter 
species became relatively abundant. Nevertheless, until the dis- 
covery tvas made, the mimicry would be an advantage to both 
species, for the reasons already adduced. In the following Table the 
colours of conspicuous insects are tabulated, i. e., those contained in 
Tables I. and IV., excluding S.fuciformis and the conspicuous larva 
of L. pini, tlie latter being omitted because I have never seen a 
specimen, and because the appearance differs greatly in the various 
figures I have been able to consult (see Table A, pp. 232-23.'i). 

I have described the colours of the images at rest to correspond 
with the larvae and puppe ; in flight tlie following effects are seen : — 
Imagos of Wasp, Bombus, JS^omada marshameU.a, E. jacobcece, A. 
filipendulce, A. grossulariatu, the tuo Coccinellidse, Telephorus and 
Chr>/some/a, would show much the same colours as at rest, although 
in E. jacobcece and A. Jilipendulce the red would be in larger amount 
because of the under wings, and in the Coccinellidse, Telep/wrus and 
Chiysomela, the black would be in far larger amount because of the 
body. The imagos of iS. menthastri, S. Jubricipeda, and P. aurijlua 
would hardly show the black spots in flight, but would appear 
whitish, yellowish, and white respectively. 

This comparison is exceedingly interesting if it is remembered that 
the colours which are repeated again and again are those which are 
known to produce the greatest effect. Thus tiie greatest {)ossihle con- 
trast is afforded by black and white, and next to this by black (or 
some very dark colour) and yellow, orange, or red, the brightest 
colours in the spectrum, which possess a far higher illuminating- 
power than any of the others. Hence we find that the colours of all 
the conspicuous insects which have been tested are in all except five 
cases included in the short list given above. And these five only 
diflf'er in the inclusion of blue in one case, and of green in the other 
four cases. Hence we probably see that in addition to the advantage 
gained by convergence which has been alluded to above, benefits have 
been derived from the colours which have been employed ; and as the 
choice of the most consjiicuous colours is limited, it is seen that a 
certain amount of similarity must follow incidentally from the 
number of forms of life among which the few combinations are 
divided. Hence convergence has been aided and perhaps given its 
starting-point by tlie action of another principle of coloration also 
favoured by natural selection, and leading in the same direction as 
convergence itself (see Table B, pp. 236, 237). 

Just as similarity in colours was favoured by the limited number 
of suitable combinations, so there are a few eminently conspicuous 



232 



MK. E. B. POIJLTUN ON TH K PROTKCTI VK [MilT. 1, 



Table A. — The Colours 



Various classes of 
Colours. 


Dark Ground-colour and Lighter Secondary Colours. 




(When iucompletely 








described in this column 














the correct details are j 




Colour nex 








given in the cohimns to Species 


Ground-colour. 


in import- 


3rd colour. 


4th colour. 




the right.) 




ance. 








I. Black and white or| Larva of V. io 


Intense black. 


White 








white and blaclt. 






points. 








2 forms. 














II. White, black, and 

yellow. 
























3 forms. 














III. Black and yellow 


Pupa of A. grossula- 


Black. 


Yellow. 








or riafa. 












yellow and black. 












10 forms. 


Imago of Xomada 
marskamella. 


Black. 


Yellow. 










Imago of Vespa \ Black. 


Yellow. 










vulgaris. 












Imago of many i Dark brown or 


Yellow or 










species of Bom- black. 
bus. j 
Larva of V. urtica Black. 


orange. 










Yellow 














points and 














often 












bands. 










Larva of P. hrassicm 


Bluish green, with. Yellow. 










black spots, so that! 










effect is veiy dark. 1 








IV. Black and red 


Imago oi E. jacobcsa 'Very dark brown ; Red. 








or 


(as seen at rest). effect black. 










red and black. 












6 forms. 


Imago of A. filipen- Greenish black ; 


Red. 










didcB (as seen at 


effect black. 












rest). 














Imago of common 


Black. 


Red. 










species of Tele- 














fhorus. 












V. Black, red, and 


Larva of D. euphor- 


Black. 


Red. 


Yellow or 






white. 


bi(B. 






white. 






4 forms. 
















Larva of P. aiiri- 


Black. 


Red. 


White. 








flua. 














Larva of E. lanes- 


Black. 


Reddish. 


White. 








tris. 














Larva of 0. atitiqua 


Dark brown ; effect 
black. 


Pink. 


White or 
yellow. 


! 





1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 233 



of Conspicuous Insects. 



Light Ground-colour and Darker Secondary Coloura. 



Species. 



Imago of S. mcnfhastri (as 
seen at rest). 



Ground-colour. 



Colour next in 
importance. 



Creamy white. 



Imago of A. grossulariata 
(as seen at rest). 

Imago of P. auriflua (as seen 

at rest). 
Larva of A. grosstdariata ... 



Imago of S. lubricipeda (as 
seen at rest). 



Larva of E. jacohcBm . 



White. 

White. 
Cream. 



Buff. 



Yellow. 



Larva of A. filipendul(B . . . Yellow, 



Larva of P. hucephala 



Black. 



Black. 

Yellow. 
Black. 



Black. 



Black. 



Black. 



Yellow (becoming orange Black, 
in parts). 



Imago of Coccinella aeptem 
■punctata. 

Imago of Coccinella bipunc- 

lata. 

Imago of Chrysomela fojpuli 



Eed. 
Eed. 
Red. 



Black. 
Black. 
Blue-black. 



3rd colour. 



4th colour. 



YeUow. 

Black. 
Orange. 



234 



MR. E. B. POULTON ON THE PROTECTIVE 



[Mar. 



Table A 



Various classes of 

Colours. 

(When iucompletelj' 

described in this cohimn. 

the correct details are 

given in the columns to 

the right). 



VI. Black, red, 
and white. 
] form. 



blue, 



VII. Brown, yellow, and 
black. 
1 form. 



VIII. Green, yellow, 
and black, or 
green, black, and 
yellow. 
4 forms. 



Dark Grouud-cohmr and Lighter Secondary Colours. 



Species. 



Larva of C. ncustria. 



Ground-colour. 



Colour next 
in import 



Larva of H. defoli- 
aria. 



Difficult to determine Orange- 
upon, but probably red. 
black, because it 
occurs so fre- 
quently between 
the other colours 
and mixed with 
them ; also under- 
side is dark. 



Reddish brown. 



Yellov 



3rd colour. 



Blue. 



4th colour. 



White. 



Black. 



and simple patterns which are in this case especially adapted for the 
respective stages of the various nauseous or dangerous insects. 

Ring-patterns. — Especially suited to the cylindrical body-form, 
such as tliat of larvae, pupee, or of images with colourless wings 
(Hymenoptera &c.). Accordingly we find this pattern developed in 
such stages, and it is also often suggested on the visible part of the 
body of other forms. 

Longitudinal Stripes.— Aho especially suited to the cylindrical 
body-form, and accordingly it is entirely found in larvae and in the 
attenuated images of the genus Telephorus. 

Spots. — Especially suited to a wide coloured expanse, such as 
that provided by the wings of Lepidoptera or the elytra of many 
Coleoptera, but also fairly adapted to the cylindrical body-form, and 
accordingly it is characteristic of conspicuous Lepidopterous and 
Coleopterous images, only two of the four included larvae possessing 



1887. J VAI.HK OF COLOUR AND MARKINGS IN INSKCTS. 2.'35 



(^continued). 



Light Ground-colour and Darker Secondary Colours. 



Species. 



Ground-colour. 



Colour nest in 
importance. 



Larva of C. verbasci 

Larva of B. c<sruleocephala 
Larva oi H. iva varia 

Larva of i). ^a^ii 



Pale green. 

Smoky green. 

Green, varying to lead- 
colour. 

Yellow, varyiug through 
light green, olive-green, 
various shades of brown 
to black (in the last 
case should be in oppo- 
site column). 



Yellow. 
Yellow. 
Y'ellow. 

Black. 



Srd colour. 



4th colour. 



Black. 
Black. 

Smoky. 

Yellow or white, 



it in at all a marked degree (A. grossulariata and A . Jilipendulce), 
and one of these is partially striped. 

Combination of Ring and Stripe, and of Ring and Spot, and Stripe 
and Spot. — Also suited to the cylindrical body-form, and occurring 
in larvae and in one imago only {E.jacobcece). 

Hence the existing arrangement of many widely separated con- 
spicuous insects possessing a similar type of pattern is due to the 
fact that there is a limited choice of available patterns, as well as to the 
factors conducing to convergence. In addition to this there is 
probably in some cases a certain amount of true mimicry in the 
acquisition of patterns and colours. Thus it is more than probable 
(as has been previously suggested by other observers) that the 
species rendered conspicuous by alternate rings of black and yellow 
gain great advantages from the justly respected appearance of Hor- 
nets and Wasps. It must not be forgotten, however, that the latter 



236 MR. E. B. POULTON ON THK PHOTECTIVE [Mar. 1, 

Table B. — The Patterns of Conspicuous Insects. 



I. Ring Patterns. 



Pupa of A. grossidariata ... 
Imago of I'espa, Komada, 
and Bomhits. 

Developed on whole length Larva of E.jacohaa 

of body. j 



Pattern developed on ab 
douiinal secmonts 



I Alternating rings of yel- 
f low and black. 






II. Longitudinal Stripes. 



Stripes numerous | Larva of P. hucepTiala 

Larva of V. urtica (a corn- 



Stripes few. 
Stripes few. 



1 



mon var.). 

Larva of P. hrassices 

Larva of D. cwndeocephala 



Stripes few Imago of TeltpJiorus. . . 

Larva of H. wavaria 
Larva of H. drfoUaria 



Smoky stripes many ; yel- 
low, one on each side. 

Stri])ed appearance only 
visible from the side. 



The one white stripe on 
each side is due to 
■white hairs ; iuur red 
stripes. 

Single dorsal while ; one 
blue, three oratige-red on 
each side, and black alter- 
nating with these in nearly 
all cases. 



Larva of P. aurifiua . 



Alternating stripes of 
black (or dark ground 
colour) and yellow. 

Smoky green and yellow 
stripes ; black dots. Per- 
haps should be included 
in VI. 

Black, with a red stripe on 
each side. 

Green (variable), with 
smoky and yellow stripes. 

Brown above and yellow 
below, narrow black be- 
tween. 

Black, with red und white 
stripes. 



Larva of C. nensfria . 



Stripes black, orange-red, 
blue and white. 



III. Spots. 



Yellow band and blotch on 
each fore wing ; spots 
numerous ; body yellow 
with black spots. 

Spots numerous 



Imago of A. grossulariata 
(as seen at rest). 



Imago of S. menthastri (as 

seen at rest). 
Imago of P. auriflua 



Larva of A. grossul-ariata... 



Few black patches and a 
yellow tuft on end of 
body. 

Orange stripe ; spots few, 
but far the more impor- 
tant feature ; otherwise 
the larva should be in- 
cluded in VI. 

Some of the spots fuse into Imago of S. luhricipeda (as 



an irregular line on each 
fore wing ; spots not nu 
merous. 



seen at rest). 



Ground. 



^^^^ite. 

White. 
Whit«. 

Cream. 
Buff. 



Spots. 



Black. 

Black. 
Black. 

Black. 
Black. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. '237 

Table B (continued). 



III. Spots {continued). 



Spots few 

Seven black spots 

Two black spots 

Spots so minute as to pro 

duee little effect. 
Spots so minute as to pro 

cluce little effect. 
Six spots 



Larva of A. filipeiidults... 
Imago of Coccinella septem- 

functcda. 
Imago of Coccinella hipunc- 

tata. 
Larva of F. to 



Ground. 



Larva of V. urticce (normal 

form). 
Imago of A. filipendu^m (as 

seen at rest). 



Yellow. 
Red. 

Red. 

Black. 

Black. 

Black. 



Spots. 



Black. 
Black. 

Black. 

White. 

Yellow. 

Red. 



IV. COMBINATIOS OF RlNQ AND StRIPK. 



Few small white spots alsO; Larva of E. lanestris. 
pre.seiit ; rings due to' 
hair-bearing tubercles. 



Black, with incomplete 
reddish rings and narrow 
white stripe on each side. 



V. Combination of Ring and Spot. 



Spots on the yellow rings. . 



Larva of C. verbasci 



Green, with incomplete 
yellow rings and black 
spots. 



VI. Combination of Stripe and Spot. 



Dots very minute and pro- 
duce little effect. 

Two spots and two stripes 
on each fore wing. 



Larva of D. euphorbia 



Imago oi E.jacob<e<e (as seen 
at rest). 



Black, with red blotches and 
stripes and white or yel- 
low dots. 

Black, with red stripes and 
spots. 



YII. Not referable to the above Types. 



One blotch on each seg 
ment. Perhaps referable 
to the spot-pattern III. 

The pink colour is scattered 
in a kind of spot and 
stripe system. 

Two unequal areas with op- 
posite colours. 



Larva of D. galii 



Larva of 0. antiqua . 



Variable ground-colour, 
with a row of black 
blotches, each containing 
a pale area. 

The effect chiefly made up 
by the dark- and light- 
coloured tufts of hair. 
Imago of Chrysomela populi Large posterior part red, 

small anterior part blue- 
black. 



The patterns in flight in the im.-jgos would be different in many cases. In E. jacoh(P(B 
and A. flipendul<s a conspicuous but confused mixture of red and black is seen. The 
slow flight of A. grossnlariata permits its pattern to be seen almost as at rest. The 
others are as above described after Table A. 

Troc. Zool. Soc— 1887, No. XVTT. 17 



238 MR. E, B. POULTON ON THE PROTECTIVE [Mar. 1, 

forms also probably gain to some extent by the greater publicity 
which follows from the resemblance. "We therefore see that the 
force which tends towards the acquisition of similar forms of colour 
and marking in widely different organisms is itself the resultant of 
other forces varying in relative amounts in the different species. 
Of these primary forces we have been enabled to detect two in the 
majority of cases, and three in the minority, i. e. (1) The feasibility 
of certain colours and patterns depending upon their effect on the 
vertebrate eye, and thus giving the enemies as easy an education as 
possible ; (2) The advantage of facilitating the education of enemies 
ijy giving them a small number of patterns and colours to learn ; 
and (3) The great additional advantage conferred by trading upon 
the reputation of a well-known and much-feared or much-disliked 
insect. 

These may probably be looked upon as the chief primary forces 
which have determined the various forms of conspicuous appearance. 
But such forces have had very different material to work upon in 
the different species, and doubtless the resultant has been largely 
influenced by the protective colours which existed before the "warning" 
colours and markings arose, and which formed the material on 
which the first steps (at any rate) were built. We can, in fact, 
point to certain conditions in the "warning" appearance of some species 
which are almost certainly remnants of a previous mode of defence 
due to protective coloration. Thus Prof. Meldola has drawn my 
attention to an opinion expressed by Mr. T. W. Wood (" Insects in 
Disguise," Student, 1868), that the larvae of ^.yorco&«« are protected 
by their resemblance to the flowers of the ragwort. I believe that 
in the orange ground-colour of this species we have such a remnant 
of a former resemblance to the flowers of this plant and the groundsel, 
on which the species also feeds. The acquisition of the black bands 
and, above all, the gregarious habits are, then, later developments 
which have followed the acquisition of an unpleasant taste. Here it 
is seen that the material at the disposal of the primary forces 
tending towards a "warning" appearance was such as to render 
most probable the resultant which has actually obtained. 

Again, Mr. Thomas Eedle informed Prof. Meldola that he believed 
the larva of C. verbasci resembles the flowers of its food-plant, 
mullein. Here, again, I entirely agree with this observation. There 
is a great deal in the larva which harmonizes extremely well with 
the yellow and dark sessile flowers, studded upon the surface of the 
thick green spike, and surrounded by green unopened flowers. In 
this case it is probable that the pattern may have been rendered a 
little more distinct ; but the very conspicuous appearance practically 
depends upon the gregarious habit, and upon the fact that the larvae 
do not chiefly rest upon the spike, but are commonly seen upon the 
uppersides of the large leaves, forming a background against which 
the larval colours appear with startling distinctness. But, as Mr. 
Eedle maintains, an isolated larva on the flowering-spike is evidently 
well protected by colour-harmony with its surroundings. These are 
but instances of the past history which must be deciphered before we 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 239 

can adequately appreciate the meaning of the colours and markings 
of any animal. 

III. Insects which evade their Enemies. 

We now come to Wallace's converse suggestion — that just as 
conspicuous forms which court observation will be avoided, so 
the insects which harmonize with their surroundings, and which 
evade their enemies, will be greedily eaten when detected and 
caught. I have adopted Meldola's suggestion that the terms 
" protective resemblance " should be applied to the appearances 
which tend to deceive enemies by their resemblance to motionless 
(vegetal or mineral) surroundings, the term "mimicry" denoting the 
resemblance to other animals. On entering upon the experimental 
investigation, I thought that I should have little to record except a 
complete agreement with everything which has been previously said 
upon the subject. I was surprised, however, to find some instances 
which are entirely antagonistic to the principles laid down by 
Wallace. Unfortunately the instances recorded by other observers 
are exceedinglj' few. Jenner Weir evidently experimented with a 
large number of species, but he gives very few details, and for the 
most part is content with summing-up his results as faA^ourable, 
without exception, to Wallace's suggestion, in these words : — " I will 
now add a few viords on those larvse which are eaten greedily by 
birds, and my remarks on the subject will be brief; it will be 
unnecessary to detail all the experiments made, as the results are 
easily generalized. 

" All caterpillars whose habits are nocturnal, dull-coloured, with 
fleshy bodies and smooth skins, are eaten with the greatest avidity. 

" Every species of green caterpillar is also much relished. 

" All Geometrse, whose larvas resemble twigs as they stand out 

from the plant on their anal prolegs, are invariably eaten 

They eat with great relish all smooth-skinned larvBe of a green or 
dull-brown colour, which are nearly always nocturnal in their habits, 
or mimic the colour or appearance of the jilant 'they frequent." 

Jenner Weir, however, gives details of experiments with other 
stages of Lepidoptera ; and I am now able to add many valuable 
details from his experiments upon Lizards in 1886. There are also 
a few instances to include from Mr. Butler's paper (already quoted) 
and a few of which I have heard from him by letter. In my experi- 
ments I chiefly made use of the imagos of Lepidoptera, as I nearly 
always sought for conspicuous larvae with which to test the sug- 
gestion previously discussed. 

Other observers having given so little detail, it follows that Wal- 
lace's converse suggestion possesses extremely little precisely recorded 
experimental foundation. There is, however, no reason to doubt 
that Jenner Weir's conclusions will be very generally confirmed by 
extended experiments, and they doubtless express the results of 
many observations. But as I have come across a few startling 
exceptions among the most protectively coloured forms, it is safer not 

17* 



240 MR. E. B. FOULTON ON THE PROTECTIVE [Mar. 1, 

to assume the existence of any great body of confirmatory evidence 
until it has all been rigidly tested and recorded. 

It will be unnecessary to separate the larvse from the other stages, 
because the meaning of imitative colours is equally clear whenever 
they occur, while the warning colours of imagos mio;ht in some cases 
be tnistaken for those of other significance. I will proceed at once 
to tabulate everything I have been able to find recorded, and vvill 
afterwards consider in detail the more remarkable cases. I have 
already imjilied that I believe the larvae o{ Papilio machaon should 
he included in the Table given below. Prof. Meldola has since shown 
me that Mr. T. W. Wood has also taken this view of tlie colours 
of P. machaon (see a paper in 'The Student,' 1868, entitled 
"Insects in Disguise"). I believe that the bright sreen colouring 
broken up by black markings is very well adapted for concealment 
among the much-divided leaves of the Umbelliferae on which the 
larva feeds. I also consider that the imago o^ S.fuciformis should 
be included (see Table V., pp. 2-l2-2.i9). 

Looking back at this list we see that as a whole its results offer 
the most decided contrast to those of the previous lists, inasmuch as 
the vast majority of species are in this case devoured with relish. 
But wliile it thus supports the converse side of Wallace's suggestion, 
this is by no means so universally true as Jenner Weir's earlier experi- 
ments led him to believe. Out of a total of 44 different species, or 
stages, of Lepidoptera, we find 7 exceptions, viz : — Imagos of S. 
ligustri, P. bucephala, and 0. antiqua ; pupae of V. to, V. urticcB, 
P. bucephala ; and the larva of il/. hjpica. Two of these appear 
for the first time in Table V., while the others have appeared before 
in other stages in the earlier tables. Deferring the consideration 
of the latter, we will take the two species rather more in detail. 

Imago of Sphinx ligustri. — I think the evidence in this case 
speaks for itself, and demonstrates very completely the protective 
importance of mere size, unaccompanied by other alarming features 
or by any means of active defence. Tlie species is admirably 
protected at rest and must be most carefully concealed. After 
twenty years, during which I have looked for insects, I have only 
once seen the moth at rest. Again, its flight is probably as rapid as 
that of any species in the world. The behaviour o( Lacerta viridis 
seemed to clearly show that the moth was highly palatable, as we 
should expect from its very perfect means of evading its enemies. And 
yet the much smaller L. muraJis would not touch the insect. If 
the supposition be raised that the moth possessed some smell, which 
was disliked by L. muralis, hnt to which L. viridis vfas indifterent, I 
can only say that I have met with no other instance of any differeuce 
of tastes when I experimented upon the two Lizards with the same 
species of insect. And if this be the correct explanation why tiie moth 
was untouched after spending many hours in the cage of L. muralis, I 
cannot but think that L. viridis would have shown some reluctance 
in devouring it, although it might have ultimately eaten it under the 
impulse of hunger. And, again, L. muralis was more ravenous than 
any of my Lizards ; and the above tables show clearly that I have 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 2 J I 

chiefly relied upon this species for the evidence that hunger will force a 
Vertebrate to eat an insect which is evidently distasteful to it. Further 
the L. viridis being less used for this purpose, were not pressed by 
hunger to the same extent as L. muralis. I think that the almost 
certain explanation is that L. muralis was afraid to touch an insect 
wliich was not far from its own size, while L. viridis was less 
timid, the difference in size being far greater. And such an 
explanation throws light on the cases already discussed, in which a 
large caterpillar is protected by gaining some marking which suggests 
the appearance of a serpent. On this jioint Weismann says (Mel- 
dola's translation as above, p. 330) : — " It does not require much 
imagination to see in such a caterpillar an alarming monster with 
fiery eves, especially if ive consider the size which it must appear 
to an enemi/ suck as a lizard or small bird." The case of »S. ligustri 
enforces this last remark, and shows how size alone may be efficient 
as a protection against the smaller insect-eating Vertebrates. So 
far as I am aware, this important use of size, unaccompanied by 
any accessory markings or any special attitude, is now brought for- 
ward for the first time ; but it is an advantage which is probably far 
from uncommon, considering tlie number of large species in the 
same position as S. ligustri, and it has doubtless been of special 
importance as an initial stage in the development of the more 
elaborated forms of terrifying appearance already described. 

Larva of Mania tupica. — In this case it was perfectly clear that 
the larva possessed a very unpleasant taste, so that it was refused by 
L. muralis even when very hungry. And yet the description given 
in the table shows that the species is highly protected in the 
larval state by protective colouring and habits which correspond. 
As the exception is so important, I will add a few details to the 
proofs given in the table. Newman gives the following facts about 
the young larv?e : when hatched from the eggs laid upon the leaves 
of pear, plum, &c., the larvae " devour the upper cuticle and paren- 
chyma of the leaf, leaving the lower cuticle entire, dry and brown ; 
they lie closely packed side by side and apparently motionless," but 
in reality gradually moving onwards, " leaving a larger brown space 
behind." We see here a most interesting adaptation of the sur- 
roundings to the brown colour of the laiva. A brown larva is 
conspicuous on a green leaf, and a single larva could not eat away 
the cuticle so as to surround itself with a brown area of sufficient 
size until after the lapse of considerable time. Hence the subordina- 
tion of gregarious habits to ends which are the exact opposite of those 
usually sought, viz. the intensification of warning colours. We 
have therefore a most elaborate and perfect mode of concealment in 
the younger stages of larval life. It is obvious, however, that such 
a metliod can only be successfully adopted wliile the larvae are very 
small, so that a large number of them can rest for a long time on a 
single leaf. Accordingly Newman tells us that "in a few days, 
perhaps from ten to fifteen, they abandon this arboreal life," and de- 
scending, feed upon many kinds of low-growing plants. It hybernates 
in October, and again feeds greedily in the following spring ; but^ 



242 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table V. — Experiments with Insects which are protectively 



Species and Stage. 



Protectiye resemblance, or habits of concealment, evasion, &c. 



Larva Saturnia carpini 



Larva Mamestra brassicce. 



1. Lepidopterous 

The green larva with its black bands and pink tubercles harmonizes remarkably 
well with the heather on which it feeds {Andrew 3Iurray : queted by Wallace 
in the essay often referred to). The larva feeds on other plants also, but the 
special relation of its appearance to that of heather seems to indicate that 
this is its ancestral food-plant. Very often, however, the larva possesses 
golden instead of pink tubercles. 



Commonest variety is olive-brown dorsally and dingy yellow ventrally, with 
abrupt line of demarcation ; a triangular mark containing two white dots on 
the back of each segment. Other varieties are brown or dingy green or any 
intermediate tint. Well concealed among the leaves or in tunnels, in 
cabbage, broccoli, &c., but freely exposed on many plants, although always 
harmonizing with the surroundings. 



Larva Tryphtena orbona... 



Larva Tryphcena pronuba . 



Colour dingy umber-brown, with darker and paler markings, 
plants, and in spring on saUow and hawthorn. 



Feeds on low 



Larva concealed by day, feeding at night on almost all the plants in gardens. 
Colour varies from pale yellowish green to dark brown, with brown, black, 
and pale markings. 



Larva Taniocampa gothica 



The whole effect of the larva is green (green ground-colour with one lateral 
white stripe, and a dorsal and two lateral very narrow pale yellow stripes). 
Hence harmonizes well with the leaves of the many plants on which it feeds. 
Disturbed it falls off and has some chance of escaping in the grass or other 
low -growing plants. 



Larva PMogophora raeticu- 
losa. 



The whole effect green or brown (for the larva is dimorphic), as the white dorsal 
and lateral stripes are inconspicuous. Hence well protected on leaves of 
food-plant, and the brown varieties on dead leaves and earth. Same protective 
habit of falling offas noticed in T. gothica. Feeds on many low-growing plants. 



Larva Mania typica The larva is coloured with various shades of brown, and is most perfectly pro- 

I tected against bro^^Ti leaves, which, as I have observed, it almost invariably 
selects, and upon which it sits motionless by day, feeding at night. If there 
are no brown leaves it retires by day into a very dark corner among the green 
leaves. It also has the habit of falling off. Feeds on many trees and low- 
growing plants. 



Larva Hyponomeuta euony- 
melius. 



Yellowish grey with black spots, not conspicuous in themselves ; but the larvae 
Uve in colonies, spinning a web, the latter certainly attracting attention. But 
the larva seem to be safe within it, as in a cocoon. The gregarious nature is 
doubtless related to the habit of spinning a common web. Feeds on spindle. 



188/.] VALUE OF COLOUR AND MARKINGS IN INSKCTS. 243 



Coloured, or which evade their Enemies by other means. 



Experimental evidence. 



E. B. Poultou. 



Larvae. 



J. Jenner Weir. 



Other observers. 



Bearing upon Wallace's 
converse suggestion. 



A. Weismann. — 
Devoured by La- 
certa viridis. 



A.G. Butler. — Eaten 
hy L. viridis. Lar- 
v£e of tbis genus and 
of Hadcna eaten by 
many birds {e.g., 
Eobins). 



Support. 



Support. 



Eaten by Lizards . 



Support, as with last species. 



Eaten by Zootoca 
vivipara. 



Strong support, from the special 
character of the coucealmeut. 



Eaten greedily by L. vmralis. 
The larvae had been found on 
Aconihim napelhcs, but the 
Lizards were unharmed by the 
poisonous food in the larval 
digestive canal. 



Eaten greedily by Hyla 



At once attacked by L. muralis, 
the larva being immediately 
detected, although rolled up 
and motionless. Nevertheless 
the Lizard evidently much 
disliked it, and after being se- 
verely bitten, it was rejected ; 
others tasted the larva with 
the same result. 



The " larvae only 
which ventured 
beyond the pro- 
tection of the web 
were eaten " by 
birds. The birds 
" appear very 
much to dislike 
the web sticking 
to their beaks." 



Support. 



Support. 



At first sight a most startling 
difficulty. Yet it was evident, 
from the behaviour of the 
Lizards, that they fully ex- 
pected the larva to be palat- 
able ; in itself a strong confir- 
mation of the suggestion that 
nearly all such larvse are 
palatable. 



Support. Being defended by the 
web there is no necessity for a 
specially perfect form of pro- 
tective resemblance. 



244 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



ABLE 



Species and Stage. 



Protective resemblance, or habits of concealment, evasion, &c. 



Pupa Fanessa to 



2. Lepidopterous 

Dimorphic : a dark grey variety with a small amount of gold, and a yellowish- 
green variety with a larger amount. Some have thought that the gold is a 
" warning" colour, but I have shown that the green form can be produced to 
the exclusion of the other by placing the larva in green surroundings before 
pupation (see Proc. Roy. Soc. vol. xlii. p. 9.3). W. H. Harwood also informs 
me that the green form is often found on the leaves of nettles, while the other 
is the ordinary form, on stones, walla, &c. Hence we have in this pupa 
the very highest kind of protective resemblance, ». e. one that is adjustable to 
the difl'erences between the various surroundings to which the organism is 
likely to be exposed. 



Pupa Vanessa uriica 



Varying from very dark grey (almost black), with hardly any gold, to very 
light pinkish varieties, with much gold, and in some cases the pupie are gilt 
all over. Iso green form. As above, the gilded appearance can be controlled 
by placing the larva in gilt surroundings (see Proc. Koy. Soc. vol. xlii. p. 95) ; 
while the dark forms may be produced by using black surroundings. 



Pupa Pygcera bucephala 



Dark reddish brown, and well protected upon or in the earth and under dead 
leaves &c., these being invariably the places chosen by the larva for pupation. 



Pupa Mamestra brassices 
(almost certainly this 
species). 



Light reddish brown : protected by being buried some slight depth in the earth. 
If accidentally exposed, it harmonizes fairly well with the earth. 



Pupa Tri/ph(Bna pronuba Reddish brown, and protected as in the last species 
(almost certainly this 
species). 



Pupa Flusia gamma ' A black chrysalis protected in a cocoon 



Imago Pieris brassicce 



3. Lepidopterous 

Protected by strong flight, and by its yellowish and black undersides harmo- 
nizing well with the yellow or white flowers of Crucifera, which it chiefly 
frequents. It is also very effectually concealed during prolonged rest (night, 
or during rain), for it is not commonly seen at rest, except in the intervals 
of flight, although such a very abundant Butterfly. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 245 



(continued). 



Experimental evidence. 



E. B. Poulton. 



J. Jenner Weir. 



Pup.«. 



Freelj eaten by L. muralis . 



Bitten by L. muralis, but evi- 
dently much disliked and 
abandoned : some were even- 
tually partially eaten when 
the Lizards were very hungry. 



Refused by the birds 



Refused by the birds 



Other observers. 



Edward Newman 
in ' British 
Moths.' — " They 
also constitute a 
favourite food of 
poultry, and are 
souglit for with 
great eagerness." 



Bearing upon Wallace's 
converse suggestion. 



A difficulty: in view of the ex- 
tremely complete form of pro- 
tective resemblance. It would 
be interesting to experiment 
with other Vertebrates and 
with hungry birds. 



Same difficulty with birds ; the 
same high form of protection. 
In this case, however, the 
Lizards freely ate the pupje, 
and they are evidently palat- 
able to these Vertebrates. 



Concerning the behaviour of the 
Lizards, it is to be noted that 
the pupal colour is not a very 
specialized form of protection, 
although complete, for it 
equally obtains in species 
which pupate in opaque 
cocoons &c. 



Eaten with great relish by L. 
muralis, slightly crushed, and 
then swallowed whole. Great 
contrast to the treatment of 
the last species. 



Edward Newman 
in ' British 

Moths.' — Fowls, 
Guinea-fowlsjPea- 
fowls, and Phea- 
sants devour them 
with the greatest 
avidity. 



Support. 



Eaten at once and evidently re- 
lished by X. muralis. 



Eaten at once (removed from the 
cocoon) by L. muralis, and 
probably relished. 



Imagines. 

Eaten readily by all Lizards, but 
not much relished, I believe, 
because of the mechanical 
difficulty of the scales and 
wmgs, and not from being 
actually unpalatable. This 
applies to all Butterflies, and 
Moths to a lees extent. 



Eaten by Lizards . . 



Roland Trimen. — A 
Swallow seen cha- 
sing this species 
(Trans. Linn. Soc. 
vol. xxvi. p. 499, 
footnote). 

A.G-.Butler.— Eaten 
by L. viridis. 



Support. 



Supj)ort. 



Support. 



246 



R. E. B. POtTLTON ON THE PROTECTIVE [Mar. 1, 



MR. E 



Table V. 



Species and Stage. 


Protective resemblance, or habita of concealment, evasion, &c. 




Imago Pieris rapce 


3. Lbpidopteuous 

Same as the last species, but not such strong flight ; on the other hand, its 
smaller size renders its concealment more perfect. 








Imago Pieris napi 


Flight rather weaker than in the last species. Protection otherwise similar, 
except that the green veining of the undersides accords well with the fact 
that the species especially prefers shady places, where green is the predo- 
minant colour. 








Imago Anthocharis carda- 
■mines. 


Flight irregular and puzzling, although not rapid. The green and white 
mottled under surface of the wings harmonizes exceedingly well with the 
green and white flower-heads of UmbelliferaB, which are especially selected 
as a resting-place (T. W. Wood, quoted by Wallace). The orange patch on 
the male's fore wing is not conspicuous on the underside. 






Flight strong: underside very dark and the insect well concealed on dark 
trunks, earth, or in shaded corners. Insect seldom seen at rest, except 
in intervals of flight, hence well concealed at night &c. 








Imago Vanessa uriicts 


As in the last species, only smaller and so more readily concealed : undersides 
not so dark and more variegated, but well concealed in the same places. 




Imago Stnerinthus populi .. 


Flight feeble : in the evening. The irregular outline of the wings, the way they 
are held, their grey and brown tints (which are alone seen in repose), all 
render the insect highly protected by suggesting withered leaves. They are 
also generally well concealed in dark corners, for they are not commonly 
seen at rest. 




Imago Sphinx ligustri 


Flight extremely powerful : in the evening. The various shades of brown of 
the fore wings and thorax are alone seen at rest, and the insect wiggests bark, 
or even more closely the wood of a cleft tree darkened with age. It is very 
rarely seen at rest, although an abundant species. 


i 


Imago Hepialua lupulinus.. 


FUght very peculiar and puzzling, consisting of rapid oscillations, always taking 
place near the ground, and for a short time at dusk and in the early morn- 
ing. At other times, when resting, very difficult to see, because of its small 
size and attitude, which exposes the brown tints of upper wings, with white 
markings, and of the body. It thus harmonizes with any of the brown 
surroundings (earth or dead leaves &e.), and is seldom detected, although 
exceedingly abundant. When disturbed it feigns death and falls to the 
ground, where it is very difficult to find. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 247 



(continued). 





Experimental evidence. 


Bearing upon Wallace's 
converse suggestion. 




E. B. Poulton. J. Jenner Weir. 


Other observers. 




Imagines {continued). 

As in the last species 


Eaten by Lizards . . . 


T.G.B., in 'Nature' 
(vol. iii. p. 166), 
has often seen 
Sparrows capture 
the species. 

A. G. Butler has seen 
the same (infor- 
mation to Prof. 
Meldola). 

A.G.Butler.— Eaten 
by L. viridis. 


Support. 




As above. Also eaten readily 
by Hi/la. 






Support. 










As above. Also eaten readily 
by Hyla. 






Support. 










Eaten readily by the Lizards, 
but not much relished, as 
above. 






Support. 










As above. Also eaten readily 
by some of the Hylce, but 
refused by others. 


Refused by all the 
Lizards. 




Support. I have no doubt that 
the refusal of some Frogs was 
due to scales only. Jenner 
Weir's Lizards were probably 
not hungry. 








Eaten by L. viridis and L. mii- 
ralis. I think more relished 
than the Butterflies. 






Support. 










Untouched by L. muralis, but 
eaten at once by L. viridis. 
Experiment repeated later 
■wiUi same result. 






Support from behaviour of L. 
viridis ; that of L. muralis is 
exceedingly interesting, • and 
probably introduces a new 
mode of protection by inti- 
midation resulting from mere 
size, with no other terrifying 
quality. 










Eaten readily by L. muralis . . . 






Support. 


\~ 







24« 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table V. 



Species and Stage. 


Protective resemblance, or habits of concealment, evasion, &c. 




Imago Cerura vinula 


3. Lepidopterous 

Flight uot powerful : in the evening. At rest the fore wings and body are seen, 
and are grey with darker markings. The insect is large and apparently not 
inconspicuous; but it must be carefully concealed, for it is uot couimouly 
found, although so abundant a species. 




Imago Pygara bucephala .. 


Flight not powerful : in the evening. At rest it is " like the broken end of a 
lichen-covered branch" {Wallace, as above). This effect is produced by the 
purple and pearly-grey colour of the fore wings, witli brown markings, and 
the ochreous tip, and by the ochreous head and tliorax. A very perfect form 
of imitative resemblance, the wings being rolled round the body so as to 
produce a cylindrical shape, with yellow ends, like a broken piece of decayed 
Btick. 




Imago Dasychira j^udibunda 


Flight not powerful : in the evening. At rest the colours, due to fore wings and 
body, are grey with darker mai-kings. Although common, it is rarely seen, 
and must be carefully concealed. 




Imago Orgyia antiqua 

(female). 


The female Moth has rudimentary wings, and never quits its cocoon, but sits 
on the outside of it, being very inconspicuous, as it is covered with grey 
down, which harmonizes well with the colour of the surface upon which it is 
resting. 




Imago Acronycta pit '. . . 


Flight rapid : at niglit. Rests by day " on the north side of trees," and there- 
fore protected by resembling lichens ( Wallace, as above). Fore wings and 
body alone seen at rest — grey, with dark markings. I have certainly often 
seen it on other aspects than north ; but anywhere on rough bark it is well 
concealed. 




Imago Mamestra brassicce .. 


Flight rapid : at night. By day well concealed by the tints of fore wings and 
part of body, which are "dark smoky grey-brown, with confused markings 
both darker and paler" {Newman); thus well protected on trunks, rocks, &c. 




Imago Mamestra persicarics 


Flight rapid : at night. By day well concealed (probably in dark corners), as it 
is seldom seen at rest. The fore wings and part of body alone .seen, and are 
" rich dark bistre-brown'' {Newman), with a white spot with a darker nucleus 
in the centre of the wing. 











188/.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 249 



{continued). 





Experimental evidence. 


Bearing upon Wallace's 
converse suggestion. 




E. B. Poulton. 


J. Jenner Weir. 


Other observers. 




Imagines {contintied). 

Eaten at once by L. viridis 






Support. 










Often refused by all the Lizards, 
with every sign of disgust, 
but they were induced to eat 
them by hunger, and seemed 
to gi't somewhat accustomed 
to the diet. Large numbers 
employed in the experi- 
ments. 






A very remarkable exception, 
when we consider the ex- 
tremely perfect protective re- 
semblance, which is so highly 
elaborated in aU its details. 


■ 








Eaten at once by Hyla, after 
other individuals had refused 
it. Offered to -L. /w((rflfo; it 
was seized and eaten directly 
it was seen to move. 






Support. 












"The only Lepido- 
pterous insect en- 
tirely rejected in 
the perfect state." 
Disregarded by all 
the birds except 
Eobin and Reed- 
Bunting, and re- 
fused by these 
after examina- 
tion. 




Another dilBciilty. 










At once taken and evidently 
much relished by L, muralis. 
Also eaten at once by some 
Frogs after refusal by others 
(experiment repeated). 






Support. 










Eaten by L muralis 


Eaten by Lizards ... 




Support. 








Eaten at once by L. muralis, 
and evidently much relished, 
as it was seized and swallowed 
immediately. 






Support. 







250 



M 



R. E, B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table V. 



Species and Stage. 



Imago TVyphsna fimbria .. 



Protective resemblance, or habits of concealment, evasion, &c. 



3. Lepidopterous 

Flight rapid : at night. By day very well concealed, probably near the ground 
and among dead bi-own leaves. Fore wings and part of body different shades of 
brown, and also olive-green in other varieties, with paler and darker markings 
and lines. Hind wings orange, with a broad black border, seen in flight, and 
with the orange and black on the undersides suggest, in the rapid motion, a 
yellow leaf blown by the wind. Probably the significance of the markings 
and the habits are the same as in the next species. 



Imago Tryph(Bna orbona ... 



Flight rapid : at night ; not so easily disturbed by day as T. prom/ha, but the 
same rapid flight and habits of concealment &c. if it is disturbed. The 
colours and protective resemblance are much the same as in the last species. 



Imago Tryphana pronuba.. 



Flight rapid : at niglit, and also easUy disturbed by day, when it flies with great 
speed, rising very quickly and dropping down (always into good shelter of 
brown leaves &c.) equally quickly. The whole process is very unlike the 
flight of a Moth, and the colour and movement suggest a yellow leaf lifted 
off the ground by a gust of wind, whirled away for a certain distance, and 
then suddenly falling agaiu ; so exact is this resemblance, that I have rarely 
been certain of the Moth until it had flown a long way. If tracked down 
and followed the Moth rises again very readily. This resemblance is chiefly 
due to the brown of the fore wings, aided by the yellow and black of the 
bind wings and the undersides. At rest it is extremely well concealed by 
the varying brown shades of the fore wings and part of the body exposed. 
It seeks dark corners, and hides deeply among thick leaves or among dead 
leaves on the ground. It is also very strong and slippery, and hard to hold 
when caught (JenJier JVeir). I have also noticed this feature. I should 
add that Jenner Weir has another theory (alluded to below) as to the 
meaning of yellow and black underwings. 



Imago Anthocelis pistacina. 



Imago Euplexia lucipara... 



Flight rapid : at night ; by day well concealed and seldom seen. The parts seen 
at rest vary much in colour, being brownish, reddish, or of different grey 
tints with faint darker markings. 



Flight rapid : at night. Very seldom seen at rest, and evidently well concealed, 
probably in dark corners and among dead leaves. Fore wings and part of 
body alone seen are rich brown with pale markings and a white mark on 
the wing. 



Imago Amphipyra pyra- 
midea. 



Flight rapid : at night ; by day well concealed and very seldom seen. The parts 
seen at rest are brown and grey-brown, with paler and darker markings. 



Imago Hadena oleracea 



Flight rapid : at night. At rest well concealed and seldom seen. Fore wings 
and part of body seen are reddish brown, with a narrow white line and two 
light spots on wings. Resembling bark or dead leaves &c. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 2.51 



(continued). 



Experimental evidence. 


Bearing upon Wallace's 
converse suggestion. 


E. B. Poulton. 


J. Jenner Weir. 


Other observers. 


Imagines (continued). 


J. Jenner Weir has 
seen a Swallow 
chase this Moth 
by day, making 
" several ineffec- 
tual attempts to 
seize it." The 
Moth, however, 
escaped. It was 
evidently consid- 
ered a very desir- 
able capture. 




Support. 








Eaten by Lizards... 




Strong support, as in T. pro- 
nuba. There is no doubt that 
it is also relished by birds. 






I have often seen it pursued by 
birds with great persistence ; 
they evidently relish it much, 
and make great efforts to 
catch it. 


Evidently much ap- 
preciated by the 
birds, but even in 
the aviary its rapid 
movements and 
slipperiness made 
it very difficult to 
capture. Eaten 
by Lizards. 




Strong support in the special 
character of the defence suited 
to so many emergencies ; pro- 
tected at rest ; again, when 
detected or disturbed, by ap- 
pearance and mode of flight ; 
and, again, when captured, 
by it slipperiness and great 
strength ; and all this coexist- 
ing with and following from 
the fact that it is keenly 
relished and much pursued. 




1 


Eaten greedily by 
Lizards, but appa- 
rently swallowed 
with difficulty, 
probably because 
of the scales. 




Support. 






1 Eaten directly by L. muralis, 
1 and evidently relished. 






Support. 








Eaten by Lizards ... 




Support. 






Eaten by L. muralis 






Support. 









252 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table V. 



Species and Stage. 



Imago Cucullia verbasci ... 



Imago Gonoptera Hbatrix .. 



Imago Ennomos angularia. 



Imago Amphidasis bctu- 
laria. 



Imago Halia wavaria 



Imago Caynjptogramma bi- 
lineata. 



Imago Chloephora prasinana 



Protective resemblance, or habits of concealment, evasion, &c. 



3. Lepidopterous 

Flight (probably) rapid: at night. At rest exceedingly well protected by a most 
joerfect resemblance, both in shape and colour, to a splinter of wood; the 
colours of the parts seen being rich umber-brown, shading into pale wainscot- 
brown, and this again into the darker colour. 



Flight rather slow : at night. At rest beautifully protected by a most special 
resemblance to a decayed red leaf with white spots of fungoid growth on it. 
The parts seen are grey with a reddish tinge, with patclies of orange-red and 
light grey lines, and two minute but intensely white spots on each wing. 
Similar white marks occur on the legs, which are often partially seen at rest. 
The effect is greatly heightened by the irregularly toothed margin of the 
wings. The Moth appears in August and September, and hybernates, so 
that it is in the perfect state when red and brown leaves are chiefly seen, and 
when green leaves are mostly absent. It selects dark places in whicli to 
hybernate — tool-houses, attics, &c. 



Flight slow : in the evening. The angulated wings are yellow with brown lines 
and short brown streaks, and this colour and shape, together with the atti- 
tude and the colonr of the undersides of wings (also often seen), all suggest 
a yellow leaf in a very perfect manner, aided by the time of appearance 
(August and September). 



Flight slow : in tlie evening. Both wings and body seen at rest, and dingy 
white with smoky markings (the latter may suffuse the whole surface in some 
viirieties). Although conspicuous when found at rest, it must generally be 
concealed with care, for it is not found very commonly at rest, although 
such an abundant Moth. The protective resemblance may be to variegated 
lichens on rocks and trees. 



Flight not rapid : in the evening ; also easily disturbed by day. Both wings 
seen at rest (as is usual in Geometrre), and they and the body are grey ; the 
wings with a purplish and brownish tinge, with brown spots and streaks 
(including the V-like mark). It is thus inconspicuous on tree-trunks &c., 
although without any special resemblance. 



Insect conspicuous on the wing. Flight not rapid : in the evening ; also very 
easily disturbed by day. Well concealed in thick leaves when at rest. Wings 
and body yellow ; former with slender white and brown waved lines. A 
variable amount of brown colouring. 



Flight rapid : in the evening. By day beautifully protected by resembling the 
green colour of foliage ; the only parts seen (fore wings and part of body) 
being yellowish green with three silvery lines across the wing ; these latter, 
on the opposite wings, come together at an angle during the attitude of rest, 
and convey the impression of leaf-veining. Moth flies in June. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 253 





{continued). 










Experimental evidence. 


Bearing upon Wallace's 
converse suggestion. 




E. B. Poulton. 


J. Jenner Weir. 


Other observers. 




Imagines {continued). 


Greedily devoured 
by the birds. 




Support. 










Eaten at once by L. vmralis, 
and evidently much relished. 
The Moth was detected while 
motionless. 




F. W. Andrewes in- 
forms me that he 
has seen a Robin 
carry the Moth 
off, having flown 
quite near to him 
iu pursuit of it. 


Strong support, because of the 
special character of protection 
and the evident keenness with 
which the Moth is pursued. 








Eaten at once by L. muralis, and 

large numbers by the Hyla ; 
although individuals would 
refuse them, yet they were 
generally taken ; and some- 
times one Frog would take as 
many as five, one after the 
other. 






Strong support, as in the last 
species. 










Eaten at once by L. muralis. 
Evidently relished. 






Support, for it certainly evades 
its enemies. 








1 




Eaten by Lizards . . . 


A. G. Butler.— 
Greedily devoured 
by Frogs. 


Support. 










Eaten by Lizards . . . 




Support, for the insect clearly 
avoids its enemies. But the 
protection does not seem to 
be very perfect. 








1 


Eaten at once and with appa- 
rent reli.sh by L. ?n>/mlis. It 
was seiEed directly it was seen 
to move, nut before. 






Strong support. 


1 








Pkoc. Zool. 


Soc— 1887, No. 


KVIII. 


18 



254 



MR. E. B. POTJLTON ON THE PROTECTIVE [Mar. 1, 



Table V. 



Species and Class or 
Order. 



Crustacea. 
Armadillo vulgaris 



Arachnid A. 
Spiders of diiFerent species. 



ProtectiTe resemblance, or habits of concealment, evasion, &c. 



4. IsrAGINES OF OTHER InSECTS AND A FEW 

(The arrangement of Claus's ' Text- 
Dark colour and habits of concealment are strongly protective. 



Epeira diadcma and Tcgenaria dmnesfica were chiefly employed in some of 
the experiments (E. B. P.), but any common Spiders which could be found 
were offered to the Lizards and Frogs. lu appearance, and especially in 
their rapid retreat into concealment, the Spiders are difficult to capture, 
especially when it has been shown that the enemies do not like the web in 
their mouths. 



Myeiapoda. 
lulus terrestris 



The dai-k colour and habits of concealment are strongly protective, but it also 
possesses a very unpleasant odour. 



Lithobius forficatus 



The brownish orange colour, and especially habits of concealment, are strongly 
protective : inodorous. 



Ortiioptera. 
Forficula auricularia 



Probably this common species of Earwig was always employed. The colour 
and habits of concealment are strongly protective. The pincers may perhaps, 
in some cases, act as " terrifying" structures. The insect has a very disagree- 
able smell. 



Periplaneta orientalis 



The colour, rapid movements, and habits of concealment are strongly protec- 
tive, but the insects also emit a very unpleasant odour. 



Becticus verucivorus. 



A large green grasshopper, with brown spots on the fore wings. 
concealed, and evidently a very powerful hopper. 



Thus well 



NeukopteeA. 
Chrysofpa, perla 



Probably this common species was employed. Its gi-een colour protects the 
insect among the leaves in which it lives, but it can also emit a peculiarly 
unpleasant odour. 






18S7.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 255 



(continued). 



Experimental evidence. 



E. B. Poulton. 



J. Jenner Weir. 



Artiiropoda of other Classes. 
book of Zoology' is followed.) 
Eaten readily by L. mitralis ... 



Always eaten with especial relish 
by the Lizards and the Frogs. 



Other observers. 



Bearing upon Wallace's 
converse suggestion. 



Spiders eaten greedily A.Gr. Butler. — Eaten 
by Lizards. by Lacerta viridis. 



Eaten readily by the Frogs. 



Eaten readily in large numbers 
by all the species of Lizards 
and by the Hyla. 



Eaten by the Lizards 



A. WeisDiann. — Ee- 
fused by i/. viridis. 



A. Weismann. — 
Greedily eaten by 
L. viridis. 



Strong support. 



Strong support. 



A modification : here are un- 
pleasant attributes coexisting 
with protective habits and 
colouring. 



Strong support. 



In this case the enemies made 
use of did not seem to object 
to the smell. Support. 



Eaten by L. viridis. 



A. Weismann. — 
Once eaten by L. 
viridis. 



As above; impleasant attributes 
coexist with protective habits 
&e., but the former do not 
protect them from tliese ene- 
mies. 



Strong support. 



Conclusion as in the case of 
Pcrij)laneta. The Lizards are 
evidently much repelled by 
certain smells, but do not 
object to others which are 
very unpleasant to man. 

1^ "~ 



256 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table V. 



Species and Class or 
Order. 


Protective resemblance, or habits of concealment, evasion, &c. 


Hemiptera. 
Aphis hedercs and Aphides, 

Bp.? 


4 Imagines of other Insects and a few Arthropoda 

AjMs hcder<B is dark and inconspicuous ; aphides are generally inconspicuous, 
but probablj also, in some cases, protected in other ways (taste or smell). 


Hcmipter07i, sp. ? 


The species made use of was inconsjjieuous ; but many possess warning-colours. 
The species was, however, evil-smelling, like the brilliant ones. 


DiPTERA. 

Muscidse of various species 


Musca domestica and M. vomitoria chiefly employed ; but also any other 
Muscida which could be found. The appearance, and especially the rapid 
flight and readiness with which they are disturbed, are strongly protective. 
Larvae and pupae also made use of : both concealed. 


Eristalis and Syr phis 


Probably the commonest species. Although somewhat less well-concealed than 
many species of Musca, many of the species of these genera are even stronger 
on the wing. 


Ti7>ul(t olcTuced 


Inconspicuous and easily disturbed 






Bibio marci 


The common male is black and inconspicuous : the much less common female 
orange-coloured and easily seen ; both fly readily when disturbed. 




COLEOPTERA. 

Mclolontha vulgaris 








Carabus hortensis (Fabr.). . . 


The dark colour and habits of concealment are certainly protective. Noc- 
turnal habits also. 


0/?iccseus Tnelunarius 


As above, the species being similar in habits and appearance, only much 
smaller. Nocturnal habits. 




Hymenoptera. 
Trichiosoma lucorum 






Cocoons of Ants, sp. ? 


Caref ullv nrotected beinff well concealed in the firalleries &c 


^1 



1SS7.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 257 



(continued). 



Experimental evidence. 



E. B. Poulton. 



J. Jenner Weir. 



OF OTHER Classes (continued). 

Aphis hedercB freely ecaten by 
young Hylm. 



Eaten greedily and in almost 
any numbers by Lizards of 
all the species and by Hyla. 
The latter was especially keen 
in capturing them, but did 
not much cure for the larvte, 
which, witli the pup;B, were 
eaten in large numbers by the 
LiEards. 



Keenly relished by the Frogs... 



Aphides? sp., eaten 
by i. viridis, L.agi- 
lis, and Z. vivipara. 
Oil another occa- 
sion hardly noticed 
by the Lizards. 



Refused by the Li- 
zards after tasting. 



Eaten witli relish by 
all the species of 
Lizards, and in 
very large num- 
bers. The larva' 
and pupre also 
eaten. 



Eaten voraciously 
by all the Lizards. 



The males eaten in large num- 
bers by young Hyke. 



Eaten by L. viridis 



Eaten by all the Lizards . 



Eaten by all the Lizards , 



Eaten bv L. mt/ra/is 



Eaten by 

viridis. 



Lacerta 



Eaten witli avidity 
by the Lizards. 



Other observers. 



Bearing upon Wallace's 
converse suggestion. 



Eaten by L. viridis 
{A. G. Butler). 



Erista lis vulpinus 
eaten in large 
numbers by L. vi- 
ridis {A. G. But- 
ler). 



Conclusion for Aphides? sp. : 
probably as above: the Lizards 
evidently di^like the taste, but 
will eat the insects when hun- 
gry. The treatment of Aphis 
hcdcreg supports conclusions. 



The unpleasant qualities evi- 
dently a defence in this case. 



Strong support. 



Strong support. 



Well known to be 
the favourite food 
of Pliea>-ants &c. 
In this case the 

species is t'ormivu 
rujli. 



Strong support. 



Support. It woidd be exceed- 
ingly interesting to compare 
the behaviour of Lizards and 
birds towards the male and 
female insects. 



Suppoi't. 
Support. 



Support. 



Support. 



Support, 



258 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 



Table V. 



Species and Class or 
Order. 



HvMENOPTERA. 

Winged females of Ants, 
sp. ? 



Workers of Ants, ap. ?. 



Apis mellifica 



Andrena n'ujro-mnca . 



Protective resemblance, or habits of concealment, evasion, &c. 



4. Imagines of other Insects and a few Artiiropoda 

Colour and some of the habits appear to be protective. The gregarious habits, 
however, make them conspicuous, but are very important iu rendering their 
acid secretion more formidable. 



As above 



Workers made use of in all cases. The brown colouring renders the insects 
somewhat inconspicuous. The comparison in this respect with the more 
formidable Wasp is interesting. 



The insects bear considerable superficial resemblance to the workers of the last 
species. 



Newman states, " always I believe on herbaceous plants, never ascen- 
ding trees." In this respect Newman is mistaken, for I have fre- 
qnently found the full-grown larva feeding on the leaves of plum in 
my own garden, and it was such an individual which was given to 
i. murahs. 

I can now add my own experience of the larval habits subsequent 
to the period at which Newman has described them. Iu the winter 
of 18S4-5, I kept a number of larvae and watched them from time 
to time throughout the wliole period of hybernation. As the room in 
which they were kept was warmed, they frequently woke up at 
night and fed upon the Calceolaria-leaves with which they were 
supplied. I was most interested in observing the extreme care with 
which they were concealed by day. If there were any brown leaves 
among the food the larvae would always get upon these, and, not 
content with the harmony between their colour and that of the leaf, 
would force their way into furrows and folds, so that they came to 
lie in deep shadow and were often quite concealed. I took some 
pfiins to see what the larvae would do when all the brown leaves were 
carefully removed, and I found that, by seeking the darkest corners 



1887.] VALUE OF COLOUK AND MARKINGS IN INSECTS. 



259 



continued). 





Experimental evidence. 


Bearing upon Wallace's 
converse suggestion. 




E. B. Poulton. 


J. Jenner Weir. 


Other observers. 




OP OTHER Classes {continued). 


Eaten by Z. vivi- 
para. 




Support; it seems clear that the 
Lizards do not much object 
to the secretion. 












Eefused by all the 
Lizards. 




It may be tliat this secretion or 
the means of using it is more 
foruiidable than in the winged 
females. 










Often seized and swallowed by 
Frogs ; but I believe nearly 
always rejected in the end, 
and very often rapidly ejec- 
ted soon after being seized, as 
though the animal liad been 
stung. Often eaten by hun- 
gry Lizards ; but tliey showed 

of seizing and disabling the 
insects. 




A.G.Butlor.— Eaten 
by Frogs, appa- 
rently with avid- 
ity. Well known 
to be eaten by 
Lizards, and that 
Spiders also catch 
the Bees easily and 
frequently. 


These facts show that even an 
insect prote,-ted by a sting 
may fall a victim to enemies 
if hungry. The comparison 
with Wasps supports Wal- 
lace's suggestion. 








Seized and swallowed by Frogs ; 
but I believe nearly always 
rejected in the end, as above. 






Conclusion as above. 









and deepest folds among the green leaves, they were nearly as well 
concealed. If a leaf became rolled up at the edge, there was certain 
to be a larva inside. The larvae were kept in a glass cylinder 
upon a plate, and the stem of the food-plant passed through a hole 
in the plate and into water in a stoneware vessel placed beneath. 
Sometimes the stem did not fit tightly in the plate, and then all the 
larvse crept through the hole and rested by day upon the stem 
above the water, where of course it was very nearly dark. I have 
had very similar experience with larvse found upon trees. I 
especially remember one instance in which the leaves were com- 
pletely removed from the young shoots on one part of a plum-tree 
trained against the wall. I could not find the larva for several 
days, but finally detected it mo&t Ciirefully concealed in the folds of 
the single brown and withered leaf which still remained on that part 
of tlie tree. 

I have now given as much information as I possess of the habits by 
which this larva renders its brown imitative colouring as efficacious as 
possible for evading the eyes of its enemies. I have gone into 
details in order to show that the larva belongs to a class which is the 



260 MR. E. B. POTJLTON ON THE PROTECTIVE [Mar. 1, 

most complete opposite of that in which the larvpe render themselves 
conspicuous in various ways. The experimental evidence shows, how- 
ever, that the larva has a most disagreeable taste and (almost certainly) 
smell, so that the most ravenous of all my Lizards would not eat it. 
It is perfectly clear that these two methods of jirotection are anta- 
gonistic if present in the degree and kind possessed by this larva. 
One of them must be useless and merely incidental, and as it is quite 
certain that the highly specialized protective colouring and habits 
of concealment are of value to the organism, the unpleasant taste 
must be the useless character. And this was seen in its treatment 
by the Lizard, for the larva was recognized at once as something 
which was expected to be palatalile, and was at first seized with great 
vigour, and it was only when the larva was injured beyond hope of 
recovery that its enemy recognized the unpleasant attributes and 
relinquished it. I witnessed the whole process ; it afforded the most 
instructive comparison with the reluctant and hesitating way in 
which a very hungry Lizard would approach a highly coloured larva 
which it knew to be distasteful. It was quite obvious that the 
Lizard fully expected a palatable insect, and was greatly surprised at 
the unwelcome result. After the larva had bled freely, another 
Lizard approached, but did not taste the insect, evidently re- 
pelled by the unpleasant smell of the freshly escaped fluids. It 
is obvious that a larva of this kind, being unpalatable, and yet giving 
off no strong smell from its surface, by which to warn its enemies, 
belonging, moreover, to an immense group of similarly protected 
insects of which the vast majority are highly relished, — it is certain 
that such a larva can gain nothing by an unpleasant taste which can 
only be appreciated after fatal injury, and which is not associated 
with any colour, marking, or habit by which tbe disagreeable 
experience could be remembered. 

We are therefore driven to the conclusion that the unpleasant 
quality is in this case a merely useless character, probably some 
incidental result of the physiological processes of digestion or 
metabolism. But such a condition is most important on theoretical 
grounds, for it at once supplies the necessary steps by which a species 
can change from one protective method to another. The most 
constant objection or difficulty which is raised against the explanation 
of the rise of any well-marked structure or function as due to the 
action of natural selection, deals especially with the initial stages. 
It is asked how natural selection can accumulate the earliest variations, 
which are (the objectors assume) of insufficient importance to act as 
criteria by which life and death can be settled. Darwin set the great 
example of giving a satisfactory answer to such objections by carefully 
working out one by one those cases in which especial difficulty was 
assumed. And here, by the instance of the larva of M. typica, we 
see at once how the difficulty of the origin of nauseous forms may 
be overcome; for this larva possesses a useless attribute ready-made 
as the incidental result of some physiological process, and at so high 
a stage of efficiency that there is no difficulty w hatever in imagining 
that it might readily oecome an important criterion of existence. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 2G1 

fallino; therefore under the influence of natural selection. Knowhig; 
that increasing efficiency in protective measures is couuterbiilanced 
by increasing keenness and cunning on the part of enemies, it is easy 
to see how, as a response to an advance l)y the latter, a species might 
take advantage of such an incidental quality to adopt an entirely 
new line of defence. The concealment of the larvse we are considering 
is evidently very successful, but if it were seen through far more 
frequently than at present, and yet the larvae were always rejected 
with disgust, there would be more and more opportunity and 
necessity for the enemies to remember the experience ; and the 
further the species varied away from the beaten path of protective 
colouring, the greater aid would it afford to memory, which, although 
that of another animal, is in this respect of far less importance for 
the possessor than for the larva itself. I need hardly point out 
that in speaking of an advance in the keenness of Vertebrate insect- 
eaters, I mean an advance in the power of detecting all such larvae, 
so that there would always remain a large proportion of palatable 
species ; while the new line of defence would only be open to such 
few of them as possess the quality of distastefulness in a marked 
degree. I am quite aware that there is another possible explanation 
of the unpleasant qualities in M. tijpica ; i. e. that they are the 
remnant of a former defence by such means accompanied by 
corresponding coloration, &c. ; but while this may exjjlain similar 
facts in the case of certain other species, I do not think that it is 
likely to hold in the instance of M. tijpica, for the protective habits 
and appearance are correlated in so perfect a manner that we are 
compelled to assume that a very long period of time must have been 
covered in the attainment of so unusual and specialized a result. 

It now remains to consider the other exceptions which are of less 
theoretical importance although of extreme interest. As the same 
species have occurred before under other tables, it will be well to 
shortly tabulate the results of all the instances among Lepidoptera 
in which experiments have been made upon more than one stage 
(see Table, p. 262). 

I much hope that future experiments will enable us to extend this 
Table, but short as it is, it appears to point to several interesting 
conclusions. In the first place there is no known instance of 
distasteful qualities in stages later than the larva when the latter is 
itself palatable. This statement will doubtless be true of the great 
majority of species however complete be theexperimental investigation, 
and it points to the conclusion that this method of defence arose 
first in the larval stage. Such a relation is to be expected ; for the 
species is exposed to more danger and is more helpless at this 
period than at either of the subsequent stages. Tlie unpleasant 
taste appeals to non-parasitic enemies which devour insects ; but the 
almost complete limitation of the attacks of insect-parasites to the 
larval stage must bear in an important way upon the other modes of 
protection in this stage, tending to produce tiiat extraordinary 
S|)ecialization in defensive methods which are well known to occur. 
The imago can escape by flight, and the pupa, if exposed, may render 



262 



MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 













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1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 2C3 

its imitative resemblance complete by entire quiescence, and it is 
usually effectually protected in other ways ; but the larva must feed, 
and at the same time is sluggish in its movements, defenceless, and 
when palatable is more relished than any other stas;?, for it does not 
possess the hard investment of tiie one or the scaly covering of the 
other. Jt has also been seen that an unpleasant taste may arise 
incidentally at this period. Assuming, then, that the great needs 
of certain larvae have been met in this way, there will be the 
tendency for the unpleasant quality to jjass on by simple continuity 
into the other stages ; and if these are hard pressed, there is always 
the possilnlity that such attributes may be made the starting-point 
of a similar method of defence for them also. Hence I believe we 
shall nearly always find that conspicuous unpalatable images develop 
from larvae which are also unpalatable and conspicuous, and such a 
conclusion is entirely borne out by tlie table. But the unpleasant 
qunlity may pass on in the same way into other stages, which hold 
their own successfully by elaborate and perfect protective resemblances, 
and then there will be no tendency for the quality to be made use 
of, although it will always remain as a possibility should the 
species be worsted by its enemies in these stages. It must be 
rem.embered that the possession of an un|)leasant taste by a 
protectively coloured species can never be injurious in any way to 
itself except in so far as it causes the destruction of a greater amount 
of insect-life, inasmuch as the part contrii)uted by the species itself 
to the total destroyed does not count as food under ordinary circum- 
stances. And the species itself remaining on the same protective 
lines as the great mass of palatable species, it will itself come in for a 
proportional share of the extra loss wliich follows from the fact that 
it is not relished as food. But so long as these unpalatable species 
remain in a small minority, the reaction of their own inedibility upon 
themselves will be inappreciable. Mr. W. Esson has kindly expressed 
the danger actually incurred in a mathematical form, showing that it 
is inappreciable when the inedible species are relatively few. If 
there were a practically unlimited number of protectively coloured 
insects consisting of two sets of species, the one set edible and the 
other inedible and consisting of individuals in the ratio of 100 : 1, it 
is reasonable to suppose that in any number n of captures there will 
be killed of each set a number of individuals proportional to the 
numbers in the sets themselves ; i. e. of the edible ^^ and of the 
inedible ^. The insect-eaters will go on catching the insects until 
the edible -^ becomes equal to the number required for their food — 
a. Therefore -^ = ^ ^^^ ^ = IW' therefore there are caught of 
the inedible species j^. that is -^. 

Considering the above-mentioned exceptions among the imagos 
rather more in detail, it would certainly be difficult to find any species 
with an appearance more completely the opposite of that produced 
by the typical warning coloration than the imagos of P. burephala 
and O. antiqua. The special character of the imitative resemblance 



264 MR. E. B. POULON ON THE PROTECTIVE [Mar. 1, 

in the former lias been previously alluded to. A friend has raised 
the ol)jectiou tliat the moth imitates a piece of twig cut cleanly at 
both ends, an object which cannot be found in nature unassisted by 
art. The reply of course is that the pur[)le and grey colour of the 
sides of the suggested cylinder, together with its pale ochreous ends 
— the one appearing to be cut transversely, the other obliquely across — 
present a most perfect resemblance to wood, with that particular 
condition of texture induced by decay, in which alone the tissue will 
break shortly and sharply as if cut, ou the application of slight 
pressure or the force of an insignificant blow. In coitu the stick- 
like appearance is admirably preserved, the two insects looking like 
a single twig (Newman). It was clear from the energetic and 
instantaneous attacks made on these moths, that tlie Lizards expected 
them to be palatable and that the unpleasant quality is incidental 
and useless. It was very interesting to observe that the coutinuity 
of the unpleasant properties through the different stages in this 
species is accompanied by a gradual lessening in their powers. The 
larva was certainly disliked far more than the other stages, and the 
pupa seemed to be more neglected than the imago, the latter being 
eaten in large numbers, although often after preliminary tasting and 
temporary rejection. This fact also favours the explanation given 
above of the origin and meaning of the unpleasant qualities in the 
two terminal stages. 

In the case of O. antiqua, we have a most inconspicuous insect 
with the same unpleasant taste. Hei'e also the same explanation 
probably holds good as far as the origin of the qualities is concerned, 
for we have previously seen that its most brightly-coloured and freely 
exposed larva was disregarded by all the birds. In this case I do 
not yet feel certain that the property may not be of some value to 
the female images ; for it would be impossible to find a more helpless 
insect, without even the power of attempting to escape by walking. 
More observations are greatly needed, and it would be especially 
interesting to ascertain whether the quickly-flyirig males are equally 
disliked as food. 

The pupa of P. bucepliala has been already alluded to. Confir- 
mation is, I think, needed of Newman's statement that fowls eat this 
pupa freely, for the dislike of the Lizards was extremely marked, and 
as a rule these animals are less delicate than birds. The pupse of V. io 
and F. urticce possess the highest form of protective resemblance ; 
and here, again, Jenner Weir's observation, I think, should be repeated, 
as the Lizards acted so very differently with the latter species. Is it 
quite certain that the birds were aware of the presence of these 
generally motionless puj)8e in Jenner Weir's experiments ? 

The consideration of the fourth subdivision of the list, including 
species of Arthropoda other than the Lepidoptera, enforces very 
strongly the conclusions of the rest of this paper, — that defensive 
habits and structures may exist in almost any kind of combination, 
so that we find plenty of instances of the co-existence of unpleasant 
attributes with protective resemblance, as well as with a " warning " 
appearance. 



I8S7.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 265 

Just as it was considered to be probable that warning-colours 
possess a sexual value for the species concerned, so it is pro- 
bable that the most extreme cases of piotective resemblance also 
have a similar significance. And in fact, when the most speci- 
alized instances of the latter kind are detected and are looked at in 
themselves, they are often seen to possess great beauty, which is 
absent from the objects they protectively resemble. To take an ex- 
treme case, the imago of Melanthia albicillata sits upon the upper- 
side of a leaf in the usual attitude of the Geometers, with its wings 
extended as if "set," and in such a position its creamy-white ground- 
colour and dark lines and blotches are very conspicuous, but most 
forcibly suggest the appearance of bird's excrement which has fallen 
on to a leaf from a great height, and has therefore been spread out into 
a large wide patch. But when the insect is detected and examined, it 
is seen to possess the greatest beauty. Thus Mr. Beauchamp says of 
it: — " The perfect insect, when bred, seems to me almost without a rival 
for purity and exquisite delicacy of design. I should doubt whether 
in the range of natural objects a more beautiful line is to be found 
than that exquisite cool grey streak upon the rich creamy ground of the 
fore wing" (Newman's ' British Moths,' p. 156). While entirely 
agreeing with this description, we should all maintain that it is very 
far from applying to the object suggested by the Moth, and which 
it nevertheless resembles very faithfully. And it is probable that 
in all cases the appearance of a sexually mature insect possesses this 
among its other meanings. 

Thus I believe that the brightly-coloured underwings of the 
genus Trypkcena have the same significance as those of Gatocala 
and of Sphincc and Smerinthus, and the same significance as the 
bright colours of the uppersides of both wing in most Butterflies, 
which are also concealed during rest. But in Tryphcena alone 
among these the bright sexually selected adornment has another 
meaning as well, and has also come under the independent action of 
natural selection. For the black and yellow colours of these 
wings, together with the colours of the undersides of both wings. 
Been during their rapid vibration in flight, greatly aid the protective 
resemblance to a dead leaf whirled along by the wind. And yet the 
very similar arrangement of red and black on the upper and under- 
sides of the underwings in Gatocala are comparatively non-protective 
and seem to have almost purely sexual significance. If, therefore, 
these brilliant colours of Gatocala were modified by natural selection 
as a response to some unusual activity on the part of its foes, if 
they became yellow and black instead of red and black, and the 
habits were correspondingly modified, we should have no reason to 
conclude that they had in consequence lost their sexual significance, 
and there is no reason for forming such a conclusion in the case of 
the genus Trypliana. 

Jenner Weir has suggested that these brightly-coloured under- 
wings have another protective meaning — that they are conspicuous, 
and hence form the mark of an enemy, and yet when seized they 
readily give way without doing harm to the insect. Again, he 



266 MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 

suggests that the enemies are startled by the sudden "manner in 
which the bright colours are displaj^ed ; but I am not aware that a 
similar significance has ever been attributed to the bright colours 
of Butterflies suddenly seen when the wings are opened. The former 
suggestion probably holds, for I think the margin of the underwings 
is more commonly found to be notched than any other part of these 
insects when captured. But the primary significance of such bright 
colours, concealed in the protective attitude of rest, must be the 
same as those of Butterflies, and I should attribute the same meaning 
— of sexual adornment — to the brilliant colours of the underwings 
of the Grasshoppers of the genus (Edipoda, also alluded to by Jenner 
Weir (see Trans. Ent. Soc. Loud. 18t)9, pt. i. p. 23). 

Conclusions. 

The following are the general conclusions arrived at by the con- 
sideration of the experimental data tabulated in this paper : — 

1. The extremely specialized defence of the larval stage follows 
from its delicate anatomical construction and the necessities which 
are imposed on it as the great feeding-stage. 

2. Highly cons])icuous insects nearly always possess some un- 
pleasant attribute, 4. e. a disagreeable taste or smell in the tissues 
and fluids of the body, or (in the case of the smell) discharged from 
special glands ; irritating hairs ; or stings. 

3. The conspicuous appearance may be due to strongly-contrasted 
colours, the presence of hairs or tufts, and the attitude in which the 
body is held, and to gregarious habits, or attention may be 
attracted by violent movements which take place when an enemy 
appears. 

4. In a small number of cases a highly conspicuous appearance 
has not yet been shown to be attended by any unpleasant attribute. 

5. In the various species in which a conspicuous appearance is 
produced by colour and marking, the same colours and patterns 
appear again and again repeated. In this way the Vertebrate 
enemies are only compelled to learn a few types of appearance, and 
the types themselves are of a kind which such enemies most easily 
learn. Furthermore certain appearances are especially impressed on 
the vertebrate foes by highly aggressive insects, feared because of 
stings &c. ; and hence there is especial advantage in any approxi- 
mation to such types. Again, the selected type of conspicuous 
appearance also depends on the (probably protective) colours which 
existed at the time when the conspicuous appearance first com- 
menced (these can be determined with a great degree of probability 
in some few cases). 

6. In a relatively few cases aggressive forms among the Vertebrata 
(Serpents) are mimicked, although such an appearance is pure 
intimidation, for the insect is quite harmless. 

7. It is not uncommon for an insect to be protectively coloured 
but when detected to assume a terrifying attitude, and in some 
cases to take up offensive measures (discharge of irritating fluid, &c.). 



18S7-] VALUE OF COLOUR AND MARKINGS IN INSECTS. 2G7 

8. A few, probably transitional, forms may be unconcealed, and 
yet not very conspicuous ; these may possess unpleasant qualities or 
may be eaten readily. 

9. The likes and dislikes of insect-eaters are purely relative, and 
if pressed with hunger the most disagreeable and highly conspicuous 
insects may be eaten. Hence probably the relatively small number 
of species which adopt such a means of defence. 

10. It seems probable that when one Vertebrate eats an unpleasant 
insect, and another refuses it, the former has conquered its prejudices, 
having originally disliked the insect. 

11. In the sexually mature forms warning colours can be distin- 
guished from sexual colours by their distribution on the surface of 
the body, by the way in which they are displayed in flight, by their 
type of pattern, and the colours employed. The sexual colours and 
patterns are beavtiful, the others conspicuous. Nevertheless, to the 
modiiied taste of a highly conspicuous insect, the warning colours 
probably possess value as sexual adornments. 

12. The conspicuous appearance has relation to the injury which 
would be inflicted by the experimental "tasting" of certain enemies, 
e. g. Birds and Lizards ; but nevertheless, other enemies, which do not 
inflict injury in tasting, e. g. Frogs, have taken advantage of the 
warning colours to a limited extent. 

13. Insects which evade their enemies by protective resemblance 
and attitude, rapid movements, or habits of concealment, &c., are 
generally palatable, but they may often possess an unpleasant taste 
or smell which may or may not protect them from enemies. 

14. In a very small number of species the most perfect form of 
protective resemblance may coexist with a most unpleasant taste. 

l.T. Mere size alone may protect a species against certain of its 
smaller foes. 

16. Comparing the diiferent stages in Lepidoptera, unpleasant 
attributes appear to arise in the larval stage, and they then often 
pass through the two other stages attended or unattended, in one or 
both, by warning colours. 

17. The most highly specialized protective colours probably also 
possess value as sexual adornment. 



Considerably over 100 species or stages of insects have been ex- 
perimented upon, and the results are described in the Tables given 
in this paper. Looking at these results as a whole, it is seen that 
the various defensive measures may exist in almost any combination, 
and that the present condition of a species is in large part an out- 
come of the means of protection in past struggles. Just as in a 
long-contested battle the same position may be taken, lost, and re- 
taken, but never held a second time with quite the same significance 
as before, because of all that has happened as a result of the previous 
occupation and of all that has happened since in other parts of the 
field, so in the ever-changing relations between a species and its 
enemies the structural and functional means of defence may be taken 



268 MR. E. B. POULTON ON THE PROTECTIVE [Mar. J , 

up, abandoned, and again taken up, but never in quite the same 
combination or with quite the same defensive meaning. 

APPENDIX I. 

J. Jenner Weir's Diaiy of Observations during 188(5. 

The Lizards with which the following experiments were made 
were : — ■ 

Lucerta viridis, two specimens, d" & ? • 
Lacerta ayilis, two specimens, c? & $ • 
Zootoca vivipara, one specimen, $ . 

Mai/ 31. — Lacerta viridis d" seized the larva of Abraxas grossti- 
lariata, and immediately dropped it, afterwards licking its jaws as if 
to remove the unpleasant taste ; the $ of the same species then 
examined the caterpillar and rejected it. 

Jutie 4. — Lacerta viridis ate the imago of a species of Chrysopa 
twice during the day ; this was the more remarkable, as these insects 
are peculiarly malodorous. 

June 9. — Lacerta viridis ate two larvae of Clisiocampa neustria, 
but afterwards refused to eat more. 

June 11. — Lacerta ayilis $ after much hesitation swallowed a 
larva of C/isiocampa neiisti-ia. 

June 12. — Larvte of C. neustria and Porthesia similis {aurijlua) 
refused by all. 

Lacerta agilis 5 ate one larva of Abraxas grossulariata, 

June 13. — The imago of Tipula oleracea eaten by 5 Lacerta 
viridis. 

June 14. — Larva of Abraxas gi-ossulariata tasted by Lacerta 
viridis cJ , and rejected. 

June 15.- — Lacerta viridis S ate Clisiocampa neustria; Lacerta 
agilis S bit the larva of Abraxas grossulariata, but refused to eat 
it, and afterwards rubbed bis nose and mouth against the moss as if 
endeavouring to remove a disagreeable taste. 

Cocoons of ants were eaten with avidity. 

Jnlg 1. — L. viridis, L. agilis, and Zootoca vivipara all ate Aphides. 

July 9. — Imagines of A. grossulariata refused by Lizards. 

One imago of Porthesia similis {aurijlua) eaten. 

Aphides scarcely noticed. 

Jw/y 31.— Imagines oHIalia wavaria and Camptogramma bilineata 
eaten. 

Spiders eaten greedily. 

August 2. — Imago of Abraxas grossidariaia refused after having 
been seized. 

August 6. — Lacerta agilis ate unwillingly, and L. viridis refused 
the larvae of Pygera bucephala. 

Zootoca vivipara ate winged 5 ant, but all the Lizards refused 
the neuters. 

August 11. — Imagines of Eristalis and of Syrphus eaten by all 
the Lizards voraciously. 



1887.] VALUE OP COLOUR AND MARKINGS IN INSECTS. 269 

August 12. — One Pi/gcera bucephala larva was eaten, but this 
species generally allowed to crawl about the cage unnoticed. 

August 27. — An evil-smelling inconspicuous Hemipteron refused 
after tasting. 

August 30. — Lizards refused to eat the gooseberry sawfly. 

August 31. — Lizards ate common earwig and imagines oi Try' 
phtBiia pronuba, T. orbona, and Ampliipyra pyramidea. 

Sept. 5. — L. viridis $ killed, but refused to swallow a humble- 
bee (Bombus). 

Sept. 14. — Lizards ate imagines of Mamestra brassicce, Pieris 
brassicce, and P. rapce. 

Sept. 27. — Zootoca vivipara ate larva of Tryplicena pronuba. 

Oct. 2. — Larvae of Tryphcena arbona eaten, and imagines of 
Anchocelis pistacina seized and eaten greedily, but apparently 
swallowed with difficulty. 

Oct. 4. — All the Lizards refused the imago of Vanessa urticcB. 

Mr. Jenner Weir also informs me that the common Muscidse were 
eaten with intense relish, their larvse and pupae being also eaten. 



APPENDIX IL 

E. B. Poulton's Diary of Observations during 1886. 

May 8. — About this date one larva of L. quercus was offered to 
L. muralis and L. viridis, but it was untouched, although allowed 
to remain many days in the cages. One imago of Pieris rapce was 
eaten. One imago of Dasychira pudibunda ( $ ) was seized and eaten 
directly it was seen to move {L. muralis). 

One larva of Mania typica was eagerly seized by two individuals 
of Z/. muralis, being detected while it was rolled up and motionless 
(feigning death). The larva was shaken and bitten, but it was not 
swallowed, and the Lizards rubbed their jaws upon the wooden floor 
of the cage, an evident sign of distaste. When the larva had been 
thus wounded another Lizard came up and inspected it closely as if 
it were going to bite, but soon retired without touching it. It seems 
probable that this last Lizard was warned by the smell of the larval 
fluids which had escaped after it had been wounded. 

Four pupae of Pygcera bucephala were introduced (Z. muralis) 
and were bitten, but at once relinquished with the signs of distaste 
described above. 

Earthworms were eaten with great avidity by all the species of 
Lizards. 

May 9. — Five imagos of P. rapce were eaten, two of them imme- 
diately. One imago of Pieris brassicce was eaten at once, being pur- 
sued by two or three Lizards {L. muralis). 

Two imagos of Pieris napi were immediately seized and eaten. 

One imago of Trichiosoma lucorum was eaten at once by L. muralis. 

May 18. — One imago of Chloephora prasinana was seized and 
eaten the instant it was seen to move but not before {L. muralis). 

May 19. — One imago of Gonoptera libntrix was at once detected 
Proc. Zool. Soc— 1887, No. XIX. 19 



270 MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 

by two Lizards even when motionless ; it was seized and eaten with 
great avidity (L, niuralis). 

One Noctua pupa (found when digging in the garden ; almost 
certainly that of Mamestra brassicce) was instantly seized, slightly 
crushed, and swallowed with great avidity {L. muralis). It was 
interesting to note the great difference between the treatmetit of this 
pupa and that of P. bucephala. 

May 21. — One imago of Anthocaris cardamines ($) was seized 
and eaten immediately (L. muralis). 

May 30. — Two imagos of C. prasinana were eaten directly with 
avidity. A few larvae of Odonestis potatoria were placed in the cage 
of L. muralis a day or two before, and after long delay had 
disappeared by this date. Two were offered to the same Lizards on 
this day, and one was taken at once, the other being imtouched for 
a long "time. It is quite evident that the species is disliked, but that 
the Lizards will eat it if they are hungry. 

Several pupae of P. bucephala were introduced ; from one of them 
an imago emerged almost immediately, and was at once seized and 
then relinquished by two or three individuals of L. muralis, but it 
was finally eaten, although evidently unpalatable. The pupae were 
not touched on this day. 

One imago of P. rapes was eaten at once. 

One imago of P. hrassicce was eaten, but not at once, by L. muralis. 
June 2. — One imago of P. rapcB was eaten at once by L. muralis. 
June 4. — One pupa of Tryphcena pronuba was eaten directly by 
L. muralis. 

One imago of Euplexia lucipara was eaten directly by L. muralis. 
Two imagos of Hepialus lupiilinus were eaten directly by L. muralis. 
One imago of P. bucephala was seized at once but soon relin- 
quished by L. muralis. 

June 6. — The P. bucephala imago introduced on June 4th had 
now disappeared ; another was offered on this day and was at once 
seized and eaten by L. muralis. 

The larvfe of O. potatoria had now disappeared. 
Three imagos of P. rupee were eaten at once by L. muralis. 
One imago of P. brassicce was eaten immediately by L. muralis. 
After this last date the various species of Lizards to which insects 
were offered were always accurately noted, and were indicated by 
the numbers L, III., IV., V., and VI, placed upon their respective 
cages, and which contained the following species : — 

I. Lacerta muralis (var. tiliguerta), about a dozen fine speci- 
mens. 
III. One $ Lacerta viridis and two d of the same species: all 

fine individuals. 
IV. L. muralis, var. tiliguerta : about half a dozen full-grown 
specimens. 
V. A few small individuals of L. muralis (more than one variety). 
VI. One 5 L. viridis, a very fine specimen, and one full-grown 
Gecko {Tarentola mauritanica). 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 271 

June 11. — I. Four imagos of P. hucephala were introduced, one 
was seized direct]}' but relinquished ; ultimately all four were eaten. 
Four larvfe of Abraxas grossulariata were untouched. 

III. Two imagos of Cerura vinula and four of P. bucephala were 
all taken ultimately. These Lizards {L. viridis) were very shy, and 
hardly ever seized an insect before an observer; being quite unlike 
L. inuralis iu this respect. 

Two larvaj of A. grossulariata disappeared, but they may have 
escaped, being small larvae. 

IV. Five larvae of A. grossulariata introduced ; I saw one severely 
bitten, in fact chewed for some time, but it was ultimately relin- 
quished. One larva of C neustria was also added, and with the A. 
rrossulariata could not be seen on the next day. It is possible that 
they may have escaped, and I do not attach importance to their 
absence, unless escape was impossible. One imago of P. bucephala 
was ultimately eaten. 

V. Two imagos of Amphydasis betularia, one of Mamestra persi- 
carice, one of M, hrassica, and one pupa of Plusia gamma were 
introduced and all eaten (I witnessed the capture of one betularia 
and the persicarice) . Four larvae of A. grossulariata were un- 
touched. 

VI. Two imagos of P. bucephala were eaten ultimately. 

June 13. — V. One imago of Acronycta psi was ultimately eaten. 

June 15. — I. Three imagos of P. bucephala were all eaten by the 
next day. 

III. Six imagos of P. bucephala were introduced, and five were 
eaten by the next day. 

V. One imago of A. curdamines ( 2 ) was eaten by the next day. 

VI. Four imagos of P. bucephala were introduced and three were 
eaten by the next day. 

In these cases the insects may have been eaten at any time between 
their introduction and the next day, when the next observation was 
taken. 

June 17. — I. One imago of Sphinx ligustri was introduced and 
untouched. 

V. One imago of Hadena oleracea and one of A. psi were eaten 
by the next day. 

June 18. — I. The S. ligustri introduced yesterday was resting on 
the upper part of the cage out of reach of the Lizards ; it was again 
placed on the floor of the case, but remained untouched. 

June 19. — III. The S. Hgustrivias still untouched in I. cage, and 
it was therefore removed and placed in III. When the next obser- 
vation was made, a few hours later, it was entirely eaten except a 
piece of one wing. 

I. Two full-fed larvae of Taniocampa gothica, found feeding upon 
Aconitum napellus, were introduced to see if they were affected as 
food by the exceedingly poisonous properties and strong taste of the 
plant upon which they had been feeding. It seemed possible that 
tlie undigested food in the larval digestive tract might be harmful to 
the Lizards, even if the insects made no further use of the properties 

19* 



272 MR. E. B. POULTON ON THE PROTECTIVE [Mar. 1, 

of their food for purposes of defence. However, the Lizards fought 
eagerly for the larvse, and the two successful ones were separated 
from the rest and remained perfectly healthy. 

June 21. — I. An imago of Smerinthus populi was eaten by the 
next day, having been seized at once. 

III. One imago of S. populi and two cockchafers (Melolontha 
vulgaris) were introduced ; by the next day the former and one cock- 
chafer had been eaten. 

June 25. — I. One imago of S. lubricipeda was eaten at once, and 
many imagos of P. hucephala. 

III. One imago of 5. ligustri was eaten in a few minutes. 

July 1. — I. One imago of S. lubricipeda and one of Macro- 
fflossa fuciformis were soon eaten, the former at once. The Lizard 
did not seize the M. fuciformis with any caution, as if afraid of a 
sting. 

July 4. — I. Three pupae and two larvae of Vanessa urticcB were 
eaten at once ; one larva of P. <turijlua was seized at once and 
chewed for some time, but it was ultimately relinquished, the Lizard 
seeming to be much irritated by the hairs, and continually opening 
its mouth. Two imagos of Ennomos angularia and one of A. psi 
were taken at once. One unnamed larva of a Sawfly was seized and 
relinquished, but apparently taken again. 

IV. Two pupae of V. urticce were soon taken. 
V. Three pupae of V. urticce were soon taken. 

VI. One imago of S. ligustri taken. 

August 14.— I. Six imagos of Vanessa io and about eight of V. 
urtices were introduced, and many were seized at once; but the 
Lizards were apparently not very eager after them, although they 
were hungry. However, in twenty-four hours all had disappeared 
except one V. io, which had got into an inaccessible place, but when 
brought down it was eaten at once. 

August 16. — I. A few larvae of V. urticce were eaten at once. 
Two larvae o( Euchelia jacobtete vvere seized at once but relinquished, 
the Lizards being very hungry. A few hours afterwards they had 
disappeared and were very probably eaten ; but I do not feel able to 
speak with confidence, as the larvae are small and might possibly 
have escaped. 

September 6. — On this date L. muralis and L. viridis were taken 
to Birmingham and offered distasteful larvae at a meeting of the 
Biological Section of the British Association. One larva of P. 
bucephaJa was placed in the cage of L. muralis, and although it was 
often very severely bitten and for some considerable time by many 
of the Lizards, it was not eaten. For a day or two before this date 
the same species of larva had been placed in the cages of i. muralis 
and L. viridis, and some of them had disappeared, so that I believed 
that they must have been eaten. Subsequently I was able to confirm 
this suspicion, for when I was removing the individuals of L. muralis 
from the travelling cage (Sept. 7), I found the faeces of one of them 
upon the floor, the excreta consisting entirely of a partially digested 
larva of P. bucephala. 



1887.] VALUE OF COLOUR AND MARKINGS IN INSECTS. 273 

I also offered (Sept. 6) the same species of Lizard a number of 
larvae of the Sawfly (Croesus septentrionalis), and although the Lizards 
seized them eagerly at first, they soon rejected them with every sign 
of disgust, the jaws being rubbed against the floor of the cage to 
remove as far as possible every trace of the unpleasant taste. How- 
ever, on the railway journey from Birmingham to Oxford (Sept. 6) 
I actually saw a hungry Lizard seize one of these larvae, and with 
much hesitation reluctantly swallow it. I was surprised at this 
behaviour, for earlier in the summer I had ceitainly seen these same 
larvae devoured with apparent avidity by nearly all the Lizards. On 
one occasion also I placed the conspicuous pupa of Abraxas grossu- 
lariata in the cage of i. muralis. I subsequently found that it had 
been bitten, and as all its contents were gone it seems certain that it 
had been at any rate partially eaten. I have also offered the imago 
of this species to the Lizards, but it has always been refused after 
tasting in some instances. Furthermore, immense numbers of pupae 
and imagos of Vanessa urticce were eaten by all the Lizards at 
various dates towards the end of August and beginning of September, 
while early in the summer humble-bees (Bombus lapidarius &c.) 
were sometimes eaten by Lacerta viridis, and the common hive-bee 
(worker) was sometimes eaten with considerable caution by most of 
the Lizards. Common wasps (queens and workers), on the other 
hand, were invariably undisturbed ; and this was also the case with 
Nomada marshamella. Cockroaches were always eaten with avidity 
by all the Lizards, as well as the common species of Muscidse, with 
their larvae and pupae. Coccinella septem-punctata was invariably 
refused without tasting. The Carabidae — Carabus hortensis and 
Omaseus melanarius — were eaten readily. The Isopod (Armadillo 
vulgaris) was also relished. 

Experiments with the Frogs {Hyla arborea, var. meridionalis) were 
less numerous and systematic ; but they yielded some very interest- 
ing results : — 

May 7, 1886. — A queen wasp was put in the aquarium, and 
immediately a Frog sprang at it and drew it into its mouth, but 
instantly recognizing (apparently by the tactile sense) the danger, 
released the insect. It is possible that the Frog was stung, but the 
whole process, capture and release, was so rapid that it is very likely 
that the animal escaped. As soon as the wasp was free a second Frog 
behaved in precisely the same manner, and after this a third. After 
this I did not see the wasp again attacked, and it was left iu the 
aquarium for twenty-four hours. 

May 9. — One imago of Pieris napi taken instantly. 

May 13. — One imago of A. carclamines ( $ ) taken instantly by 
one Frog after being refused by others. 

May 29. — One imago of A. cardamines ( $ ) eaten at once by one 
Frog after having been refused by others. 

Oue imago of Orgyia pudibunda {<S) eaten at once by one Frog 
after having been refused by others. 

June 6. — Two imagos of E. jacobcea. were eaten at once, one 
directly after the other, by the same Frog, so that the taste could not 



274 MR. G. A. BOULENGER ON FISHES COLLECTED [Mar. 1, 

have been unpleasant. However, they were evidently indigestible, 
for next day both were found floating in the aquarium. 

One larva of A, grossulariata was refused after being just tasted by 
one Frog. 

These are all the regular notes made upon the insects eaten by 
the Frogs, but in addition to the above various other larvae and imagos 
were given to them. The foUowingimagos were eaten : — E. angularia, 
V. urticce (both these in great numbers, although they were often 
refused by individual frogs), Acronycta psi. 

The following larvae were also eaten : — Phlogophora meticulosa, 
and the hymenopterous Crcesus septentrionalis. 

Although wasps were refused, the common hive-bee was eaten, 
together with other species of bees (e. g. Andrena nigro-cenea) and 
many species of Diptera {e. g. common species of Miisca, Eristalis, 
and Syrphus, Bibio mnrci, &c.) and of spiders (e.g. Epeira diadema, 
Tegenaria domestica, &c.). All of these were relished and eagerly 
sought after except the bees, which were generally swallowed, but in 
most cases rejected afterwards and were found floating in the aqua- 
rium. Very often I saw the bees (Apis and Andrena) liberated after 
being held in a Frog's mouth for some seconds, and as soon as the 
animal began to reject it most violent and active efforts were made, 
especially with the tongue, in order to get rid of the insect as rapidly 
as possibly. From the sudden and spontaneous way in which the 
insect was often rejected after being held in the mouth for some 
seconds, I was led to believe that the Frog was stung. Earthworms 
were eaten by some of the Frogs, but apparently without relish, and 
the majority refused them altogether, and the same was true of the 
larvae of the commonest Muscidse. Goccinella septem-pitnctata and 
C. bipuncfata were invariably refused. Cockroaches, Earwigs, and 
Aphis hedercE were eaten, the latter by very young Frogs, 



2. An Account of the Fishes collected by Mr. C. Buckley in 
Eastern Ecuador. By G. A. BoulengeRj F.Z.S. 

[Eeceived February 7, 1887.] 

(Plates XX.-XXIV.) 

The rich collections brought over from Ecuador by the late Mr. 
Clarence Buckley in 1880 contained a large number of highly inter- 
esting and well-preserved Fishes obtained at three localities, viz. 
Canelos, Sarayacu, and Pallatanga. On the arrival of the collection 
a set of all the species was selected and retained for the National 
Museum. The duplicates having been sold by the well-known 
dealer Mr. Gerrard to other institutions, principally to the Vienna 
Museum, some of the novelties have already been described by Ur. 
Steindachner. 



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1887.] by mr. c. buckley in eastern ecuador. 275 

Chromides. 

1. AcARA sYSPiLus, Cope. 

Acara syspilus. Cope, Proc. Acad. Philad. 1872, p. 2.55. 
Canelos. 

2. Crenicichla saxatilis, L. 

Canelos. 

SiLURIDJi. 

3. PiMELODUs BUCKLEYi, sp. n. (Plate XX. fig. 1.) 
Pimelodus lateristriga (non Miill. & Trosch.), Cope, I. c. p. 270. 

D. 1/6. A. 12. P. 1/9. 

Near P. lateristriga. Head naked above ; occipital process 
narrow, thrice as long as broad, extending to the basal bone of the 
dorsal spine. Adipose fin much developed, a little more than one 
third of the total length (without caudal) ; its distance from the 
dorsal fin is less than the length of the latter. The maxillary 
barbels extend to the origin of the anal, the outer mandibulars to 
the extremity of the pectorals. The length of the head is one fifth 
of the total length (without caudal) ; eyes of moderate size, occupy- 
ing the middle of the head. Dorsal fin much higher than long; 
the spine much shorter than the anterior branched rays, two thirds 
or three fifths the length of the head. Pectoral spine rather longer 
than dorsal spine, smooth on its inner edge, feebly serrated at the 
extremity of its outer edge. Caudal fin deeply cleft, with the lobes 
pointed, the upper being the longer. A dark brown spot on the 
shoulder, at the origin of the lateral line ; a blackish streak along 
the latter ; upper half of dorsal blackish; adipose fin with a fine 
dark brown edge. 

Total length 150 milHm. 

Two specimens from Canelos. 

4. Pimelodus (Rhamdia) longicauda, sp. n. (Plate XX. 
fig. 2.) 

D. 1/6. A. 10. P. 1/9. 
Head naked above ; occipital process short, widely separated from 
the dorsal spine. Adipose fin long, its length being contained once 
and one third to once and three fourths in the total (without caudal) ; 
its distance from the dorsal fin is one half or three fifths of its own 
length. The maxillary barbels extend to the base of the ventrals, 
the outer mandibulars to the axilla. The length of the head is one 
sixth of the total length (without caudal) ; the depth of the body 
below the dorsal equals the depth of the tail above the anal, and is 
contained nine or ten times in the total length (without caudal). 
The lower jaw is the shorter ; the band of praemaxillary teeth is about 
four times as broad as long. Eye equally distant from the end of the 
snout and the gill-opening ; its diameter a little less than the width 
of the interorbital space. Dorsal fin higher than long, with the 
spine very feeble. Pectoral spine very feeble, not striated. The 



276 MR. G. A. BOULENGER ON FISHES COLLECTED [Mar. 1, 

posterior anal rays do not extend to the vertical from the end of the 
adipose fin if laid backwards. The free portion of the tail is as deep 
as long. Caudal fin deeply forked, the upper lobe much produced, 
much longer than the lower, measuring more than one fourth of the 
total length. Upper parts brownish, lower whitish. 

Total length 1/5 millim. 

Four specimens from Canelos. 

5. PiMELODUS (PSEUDOPIMELODUS) PULCHER, Sp. n. (Plate 

XXI. fig. 1.) 

D. 1/6. A. 9. P. 1/5-6. 

Head naked above ; occipital process short, about as long as and 
in contact with the basal bone of the dorsal spine. The length of 
the adipose fin equals about three fourths of its distance from the 
dorsal, or the depth of the tail below its origin. The maxillary 
barbels extend to the base of the pectoral spine ; the outer mandi- 
bulars a little shorter than the maxillaries. The length of the head 
is about two sevenths of the total (without caudal) ; the depth of 
the body below the dorsal one fifth or one ninth of the total length 
(without caudal). Head slightly longer than broad. The band of 
teeth in the upper jaw is of moderate breadth, without prolonged 
lateral portion. Byes very small, directed upwards, and covered 
with skin. Dorsal fin a little higher than long, with strong serrated 
spine. Pectoral spine very stout, depressed, very strongly serrated 
along its inner, less so along its outer edge. Caudal forked. Pale 
brownish on the head and body, with a dark brown band encircling 
the body and covering the dorsal fin, save its upper border, which is 
white ; head dotted with brown ; tail and caudal dark brown, with 
a large round light spot on each side of the free portion of the tail 
(sometimes confluent) ; two large whitish spots, one above the other, 
on the caudal, the extremity of which is whitish ; adipose fin dark 
brown, whitish in front and behind ; pectoral and ventral with one, 
anal with two dark brown cross bands. 

Total length S7 millim. 

Three specimens from Canelos. 

6. Cetopsis plumbeus, Stdr. 

Cetopsis plumbeus, Steind. Denkschr. Ak. Wien, xlvi. 1883, p. 31, 
pi. vi. fig. 3. 
Sarayacu. 

7. Stygogenes humboldti, Gthr. (Plate XXI. fig. 2.) 

One specimen, 56 millim. long, from Pallatanga. Specimens from 
Canelos are mentioned by Steindachner. They are of great interest 
as settling the point of the exact habitat of the species, those upon 
which it was established being without locality. Whether S. hum- 
boldti is identical with Humboldt's Pimelodus cyclopum must remain 
an open question. The opinion of Putnam (Amer. Nat. 1871, p. 694) 
loses all value from the fact that he also proposes to unite Arges 
brachycephalus, Gthr. ! On comparison of young specimens of the 



1887.] BY MR. C. BUCKLEY IN EASTERN ECUADOR. 277 

latter species with others obtained by Mr. Edward Whymper at 
Miiligalli, Ecuador, and which are undoubtedly the Brontes prena- 
dilla, C. & v., I am convinced that Steindachner's recent suggestion 
that A. brachi/cephalus is identical with A. prenadilla is correct. 

8. Plecostomus bicirrhosus, Gron. 
Two young specimens from Canelos. 

9. Ch^TOSTOMUS CIRRHOSUS, Val. 

Canelos. 

10. Ch^tostomus microps, Gthr. 
Canelos. 

11. Ch^tostomus DERMORHYNCHUS, sp. n. (Plate XXII.) 

D. 1/8. A. 1/5. P. 1/6. V. 1/5. L. lat. 25. 

Allied to C. microps, Gthr., and C. nudirostris, Ltk. Head and 
body much depressed, without any prominence ; the width of the 
head equals its length, and is one third of the total (without caudal) ; 
the entire margin of the snout naked, soft, swollen, without tentacles ; 
fold of the upper lip short, not prolonged in the middle ; barbel 
very short. Diameter of the eye about one sixth of the length of 
the head, and three fifths of the width of the interorbital space. 
Interoperculum with four or five strong, hooked, erectile spines, 
none of which are as long as the diameter of the eye. Thorax and 
abdomen entirely naked. Dorsal fin slightly higher than long ; the 
length of the anterior rays nearly equals the length of the head ; the 
basal length of the fin is less than its distance from the caudal ; six 
scutes between the two dorsal fins. Caudal fin feebly emarginate, 
lower lobe longest ; anal moderate ; ten scutes between anal and 
caudal. Pectoral spine of moderate strength, extending beyond the 
root of the ventral. Scutes of body not keeled, with short spines 
along the margin ; posthumeral ridge indistinct. Olive-brown above, 
each dorsal scute with a rather indistinct light central dot ; an 
indistinct dark lateral band ; dorsal fin with light dots ; lower 
surfaces whitish. 

Total length 148 millim. 

Three specimens from Canelos. 

12. LORICARIA FILAMENTOSA, Stdf. 

Loricaria filamentosa, Steind. Denkschr. Ak. Wien, xxxix. 1879, 
p. 45, pi. ix. 

Canelos. 

13. Loricaria lanceolata, Gthr. 

Loricaria lanceolata, Giinth. Proc. Zool. Soc. 1868, p. 235, fig. 3. 

Two specimens from Canelos, one of which agrees in every respect 

with the type. The other, an adult male with long hair-like bristles 



278 MR. G. A. BOULENGER ON FISHES COLLECTED [Mar. 1, 

on the sides of the snout, the nape, and the pectoral fin, differs in 
the much smaller size of the pectorals, which do not reach the base 
of the ventrals. I must add that the ventral and dorsal scutes of 
the three specimens before me agree perfectly with the accurate 
figure of L. magdalenee, Stdr. (Denkschr. Ak. Wien, xxxix. p. 74, 
and xli. p. 26, pi. vii. fig. 2). 

14. ACESTRA KNERI, Stdr. 

Acesti-a Tcnerii, Steind. Denkschr. Ak. Wien, xlvi. 1883, p. 26, 
pi. vii. fig. 1. 

Cauelos and Sarayacu. 

15. BUNOCEPHALUS KNERI, Stdr. 

Bunocephalus hnerii, Steind. I. c. p. 9, pi. ii. fig. 2. 
Canelos. 

16. Trichomycterus kneri, Stdr. 

Trichomycterus knerii, Steind. Sitzungsb. Ak. Wien, Ixxxvi. i. 
1882, p. 81, pi. V. fig. 1. 
Canelos. 

Nannoglanis, g. n. (Trichomycterina). 

Adipose fin present, large. Dorsal fin short, without pungent 
spine, placed nearly in the middle of the body and behind the 
ventrals fins ; anal short ; caudal truncated. Teeth viliiform, in 
broad bands in the jaws ; palate toothless ; cleft of the mouth 
moderate. No nasal barbel; one maxillary and two lateral mentals. 
Eyes directed upwards. Head covered with soft skin. No opercular 
or interopercular armature. Gill-openings wide, continuous across 
the throat. Ventrals six-rayed. 

17. Nannoglanis fasciatus, sp. n. (Plate XXI. fig. 3.) 

D. 7. A. 8. P. 9. V. 6. 

The length of the head is one fifth of the total, the height of the 
body one ninth. The diameter of the eye is one third of the length 
of the snout and three fourths of the width of the interorbital space. 
The maxillary barbel extends to the middle of the pectoral, the outer 
mandibular not quite so far. The origin of the dorsal is in the 
middle between the end of the snout and the extremity of the adi- 
pose fin, which is as long as the head. Pectorals not quite reaching 
the base of the ventrals. Vent situated below the origin of the 
dorsal fin. Yellowish, with four broad brown, black-edged cross 
bands above ; the first is the broadest, and occupies the space 
between pectorals and ventrals ; the third is below the adipose fin ; 
and the fourth, narrowest, at the base of the caudal ; a dark brown 
line from the eye to the maxillary barbel. 

Total length 52 millim. 

Two specimens, without particular locality. 



1887.] BY MR. C. BUCKLEY IN EASTERN ECTJADOR. 279 

18. Stegophilus punctatus, sp. n. (Plate XXI. fig. 4.) 

D. 8. A. 7. P. 6. V. 5. 

Closely allied to S. macrops, Stdr. Head as long as broad ; its 
length is contained six and a half times in the total, the depth of the 
body nearly nine times. Eye large, covered with skin ; its diameter 
equals the length of the snout and is contained four times in the 
length of the head. Barbel shorter than the eye. Anal behind 
the dorsal, the origin of which is nearly midway between the occiput 
and the extremity of the caudal. Latter fin emarginate. Pale brown 
above, with numerous small brown spots ; a lateral series of large 
rounded purplish-brown spots ; dorsal and caudal brown-spotted. 

Total length 1 14 millim. 

Cauelos. A single specimen. 

ChARACINIDjE. 

19. Curimatus dobula, Gthr. 

Curimatus dobula, Giiuth. Proc. Zool. Soc. 1868, p. 243. 
Curimatus nasus, Steind. Sitzungsb. Ak. Wieu, Ixxxvi. i. 1882, 
p. 80, pi. V. fig. 2. 
Canelos. 

20. Parodon buckleyi, sp. n. (Plate XXIII. fig. 1.) 

D. 12. A. 9. P. 17. V. 8. L. lat. 37. L. transv. 9. 

Dental formula -gug- ; prsemaxillary teeth fringed rather than 
denticulated, each with about twenty fringes. The height of the 
body is not quite one fourth of the total length (without caudal), 
the length of the head one fifth. The height of the dorsal a little 
exceeds the length of the head ; its origin is nearer the adipose fin 
than the end of the snout, and falls above the thirteenth scale of the 
lateral line. A length of six scales separates the extremity of the 
pectoral from the base of the ventral, which falls below the middle 
of the dorsal ; ventrals extending slightly beyond the vent. Upper 
half pale brownish, lower yellowish, separated by a greyish band ; a 
brown band along each side of the back ; fins unspotted. 

Total length 13.5 millim. 

A single specimen from Canelos. 

This being the first specimen of the genus Parodon received by 
the British Museum, the characters enumerated in the following 
tabular synopsis of the species hitherto described are merely the 
result of compilation. The shape of the praemaxillary teeth of 
P. bucTtleyi is clearly quite distinct from that of the species estab- 
lished by Kner and by Reinhardt, who describe and figure each 
tooth with about ten or twelve denticles. Whether the new species 
differs in this respect from the type of the genus 1 am not able to 
say, Valenciennes's figure not being executed with sufficient accuracy, 
and the description merely stating " le bord (des dents) est denticule 
et comme finement frange." 



280 



MR. G. A. BOULENGER ON FISHES COLLECTED [Mar. 1, 



1. P. stiborbiialis, C. & V. 

2. P. bucJdeyi, Blgr. 

3. P. nasus, Kner..., 

4. P. hilarii, Ehdt. 

5. P.affinis, Stdr... 



J). 


A. 


P. 


L. lat. 


Maud, 
teeth. 


11 


9 


16 


37 


3-3 


12 


9 


16 


37 


3-3 


11 


9 


15 


36-38 


3-3 


11 


9 


15 


38 


2—2 


12 


8 


12 


44-45 


2—2 



Height of body 
contained in 
total length. 



More than six 

times. 
Less than six 

times. 



21. Characidium fasciatum, Rhdt. 

Characidium fasciatum, Reinh. Overs. Vidensk. Forh. 1866, p. 56, 
pi. ii. figs. 1, 2. 

D. 11. A. 8. V. 9. L. lat. 36-37. L. transv. A. 

The height of the hody equals the length of the head, and is 
contained four times and one third to four times and three fifths in 
the total length (without caudal). Nasal openings widely separated 
from each other, as in the types (one of which, presented by Prof. 
Reinhardt, is in the British Museum)'. Suboperculum rather 
strongly produced and angular posteriorly. Snout and eye equal in 
length, measuring about one fourth the length of the head. Origin 
of the dorsal a little nearer the adipose fin than the end of the snout. 
Pectorals extending to the base of the ventrals, which do not reach 
the anal. Brownish (probably hyaline in life), with a broad silvery 
band along the lateral line, and more or less distinct traces of ten 
or eleven dark transverse bands on the body and tail ; a purplish 
band across the base of the six posterior dorsal rays, a small round 
blackish spot on the base of the caudal, at the termination of the 
silvery lateral band. 

Total length 87 millim. 

Four specimens from Sarayacu. 

Dr. Steindachner (Sitzungsb. Ak. "Wien, Ixxxvi. 1882, p. 78) men- 
tions C. fasciatum from Canelos, and describes, from the same locality, 
a new species, C. lourpuratum (C etheostoma. Cope ?), of which I 
am sorry to find no specimens. 

22. Leporinus striatus, Kner. 
Canelos. 

23. PiABuciNA elongata, sp. xi. (Plate XXIII. fig. 2.) 

Piabucina uniteeniata (non Giinth.), Steind. Denkschr. Ak. Wien, 
xlvi. 1883, p. 41. 

D. 10. A. 12. V. 8. L. lat. 30. L. transv. 8. 

^ Steindachner's statement in his description of C. fasciatum (Sitzungsb. Ak. 
Wien, Ixxiv. 1877, p. 559), " Entfernung der Narinen von einander gering," is 
somewhat surprising. 



1887.] BY MR. C. BUCKLEY IN EASTERN ECUADOR. 281 

The height of the body is considerably less than the length of the 
head, and is one fifth of the total (without caudal) ; the length of 
the head is contained about four times and one fourth in the total 
(without caudal). Lower jaw obtuse, projecting beyond the upper ; 
the inner borders of the mandibles closely approximate anteriorly, 
diverging posteriorly, the part of the chin exposed between them 
being ^-shaped, as in P. unilaniata \ The maxillary extends 
beyond the anterior margin of the orbit. The diameter of the eye 
is nearly half the width of the interorbital space, a little less than 
the extent of the snout, and one fifth of the length of the head. 
The origin of the dorsal fin is nearer to the root of the caudal than 
to the end of the snout, and behind the vertical from the base of the 
ventral. Adipose fin very small ; caudal forked, with its basal half 
scaly. The length of the pectoral is two thirds of that of the head, 
and exactly one half of its distance from the ventral. Ventral shorter 
than pectoral. Pale brown above, yellowish inferiorly ; a black 
lateral band becoming greyish and rather indistinct in the adult; 
a black spot on the base of the anterior dorsal rays, another on the 
root of the caudal fin. 

Total length 145 millim. 

Two adult specimens from Canelos, and three young from 
Sarayacu. 

24. Tetragonopterus rutilus, Jen. 
letragonopterus fasciatus, Gthr. 
Canelos. 

25. Creagrutus muelleri, Gthr. 
Canelos. 

26. Paragoniates albtjrnus, Stdr. 

Paragoniates alburnus, Steind. Sitzungsb. Ak. "Wien, Ixxiv. i. 
1876, p. 117, pi. viii. fig. 3. 
Canelos, 

Leptagoniates, g. n. 

Body elongate, very strongly compressed. Dorsal fin short, 
placed behind the middle of the length of the body, far behind the 
ventrals ; anal very long, nearly two thirds the length of the body. 
Cleft of the mouth narrow ; prsemaxillary, maxillary, and mandible 
with a single series of tricuspid teeth. Gill-openings wide. Scales 
moderate. Lateral line complete. 

The nearest ally of this new genus is Paragoniates, Steind., which 
differs in the following points : — Cleft of the mouth wide ; anal 
originating very slightly in advance of the dorsal ; lateral line 
interrupted. 

' In P. erythrinoides, 0. & V., the inner borders of the mandibles are widely 
separated in front and nearly parallel. 



282 MR. BOULENGER ON FISHES FROM ECUADOR. [Mar. 1, 

27. Leptagoniates steindachneri, sp. n. (Plate XXIII. 
fig. 3.) 

D. 10. A. 70. V. 8. P. 12. L. lat. 47. L. transv. |. 

The deptb of the body is one fourth of the total length (without 
caudal), the length of the head one sixth. Mandible strongly pro- 
jecting beyond the mouth ; maxillary not reaching below the anterior 
border of the eye; praemaxillary teeth 1.5, maxillary (on each side) 
11, mandibular 14; mandibular teeth largest, maxillary smallest. 
The diameter of the eye equals nearly two fifths the length of the 
head, and exceeds the width of the interorbital space. The pectoral 
fins reach nearly the extremity of the veutrals, which are small ; 
the dorsal originates above the 23rd anal ray. Colourless ; sides of 
head and a lateral band above the lateral liue silvery. 

Total length 95 millim. 

A single specimen from Sarayacu. 

28. Anacyrtus pauciradiatus, Gthr. 
Canelos. 

29. Anacyrtus knerii, Stdr. 

Anacyrtus knerii, Steind. Denkschr. Ak. Wien, xxxix. 1879, 
p. 65. 

Cynopotamus humeralis, Kner, nee Val. 

Canelos. 

GYMNOTIDiE. 

30. Sternarchus albifrons, L. 

Canelos. 

31. Sternarchus (Rhamphosternarchus) curvirostris, 
sp. n. (Plate XXIV.) 

Snout produced into a long narrow tube, which is bent down- 
wards ; the diameter of this tube, halfway between its extremity 
and the eye, is one eighth of the length of the snout. The 
distance between the eye and the base of the pectoral equals 
two thirds the length of the snout. INIouth very narrow, its cleft 
not twice as long as the diameter of the eye. Vent below the eye. 
Anal commencing nearer the eye than the gill-opening; 185-188 
rays. The greatest depth of the body is contained once and three 
fifths in the length of the head, and five times in the total. Scales 
on the upper and lower parts very small, those in the middle of the 
side of moderate size. Uniform brown. 

Total length 125 millim. 

Two specimens from Canelos. 

32. Sternopygus carapus, L. 
Canelos. 

33. Carapus fasctatus. Pall. 
No particular locality. 



1887.] ON A VESTIGIAL STRUCTURE IN THE OSTRICH. 2>^3 

EXPLANATION OF THE PLATES. 

Plate XX. 
Fig. 1 . Pimelodiis buckleyi, p. 275. 

2. Pimelodiis {Ehamdia) lo7igicauda, p. 275. 

Plate XXI. 

Fig. 1. Pbnehdus (Pseudopimelodiis) pulcher, p. 276. 

2. Sfygogenes humholdti, p. 276. 

3. Nannoglanis fasciatus, p. 278. 

4. Stegophilus punctatus, p. 279. 

Plate XXII. 
ChcBtostomus dermorhynchut, p. 277. 

Plate XXIII. 
Fig. 1. Parodon buckleyi, p. 279. 

2. Picdmcina elongata, p. 280. 

3. Leptagoniates steindachneri, p. 282. 

Plate XXIV. 
Sternarchiu {Ehamphosternarchus) curvirostrie, p. 282. 



3. Note on a Vestigial Structure in the Adult Ostrich repre- 
senting the Distal Phalanges of Digit in. By Richabd 
S. Wray, B.Sc. (Communicated by Professor Flower.) 

[Eeceived February 2, 1887.] 

While examining an Ostrich's wing in the fresh state in order to 
make out the relation of the quill-feathers to the bones, I was struck 
by observing that the phalanx of the third digit had a large amount of 
cartilage at its tip. Having another wing available with that part un- 
injured, I removed the skin covering it, and carefully dissected out the 
phalanx of digit in. From the tip of this there extends a round 
band or rod of cartilage about half the length of the first phalanx ; 
at its base it is as broad as the tip of the phalanx, at the other end 
about one sixteenth of an inch in breadth. Its distal end fades into 
the connective tissue in that region. When first dissected out, the 
cartilaginous rod showed no signs of ossification ; but when placed 
in glycerine, the rod became quite transparent, showing a free second 
phalanx embedded in it, and occupying its proximal third (see fie. 1. 
p. 284). rj o f \ & y 

The shape of the phalanx of digit in. is often as shown in fig 2, 
the pointed end being the fused second phalanx, which in adult spe- 
cimens may be free and embedded in cartilage. The adult Ostrich, 
therefore, presents the nearest approach to the pentadactyle manus 
among Birds. 

Through the kindness of Mr. G. B. Howes, I have had the oppor- 
tunity of examining some fore limbs of Ostrich embryos. These (see 
fig. 3) show a most interesting appearance, the outline of the digits is 



284 



MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 



clearly seen where they are encased in the skin. The tips of all the 
digits are free, including digit til, which has its tip free and projec- 
ting beyond the wing-fold. On removing the skin and examining 
the skeleton, phalanx 1 is distinct, then a rod of cartilage extends 
to the tip of the projecting fold of skin (a, fig. 4). This rod of carti- 




Fig. 1. Phalanx 1 and the vestigial cartilage of digit iii., adult Ostrich. Ph. 1, 
1st phalanx ; Ph. 2, 2nd phalanx ; r, vestigial cartilaginous rod ; c, 
connective tissue. 

Pig. 2. Phalanx of digit iii. of another adult Ostrich, showing Ph. 2 ankylosed. 

Fig. 3. The distal part of digit iii. in the manus of the embryo (fig. 4). 

Fig. 4. Ventral view of left manus of embryo, a, free tip of digit iii. 

lage probably represents the remaining phalanges of the digit, which 
are never definitely difi'erentiated except phalanx 2 : this is ossified 
in the broader basal third of the rod ; in the embryo, before ossifi- 
cation commences, the basal part is much the broadest. All this 
points to the conclusion that this cartilaginous rod is a vestigial 
structure, representing in addition to the second (ossified in the adult), 
the third probably, and possibly also the fourth, phalanx of digit iii. 
(see figures 1 and 3). 



4. On the Terrestrial Mollusks of the Viti Islands. — Part II.' 
By Andrew Garrett^ of Huahine^ Society Islands. 
(Communicated by Mr. John H. Ponsonby, F.Z.S.) 

[Eeceived December 8, 1886.] 
Genus Melampus, Montfort. 

1. Melampus luteus, Quoy & Gaimard. 

Auricula lutea, Quoy & Gaimard, Voy. Astrol. ii. p. 163, pi. 6. 
figs. 25-27 ; Deshayes, Lam. Hist. viii. p. 388 ; Kuster, Auric, 
p. 39, pi. 6. figs. 1-3 ; Mousson, Jav. Moll. p. 47, pi. 5. fig. 6. 

1 See Part I., siqyra, p. 164. 



1887.] MOLLUSKS OF THE VITI ISLANDS. 285 

Conovulus luteus, Anton, Verz. p. 48. 

Melampus luteus. Beck, Ind. p. lOG ; M. E. Gray, Figs. Moll. 
Anim. pi. 306. fig. 5; H. & A.Adams, Proc. Zool. Soc. 18.54, 
p. 10; Gen. Moll. ii. p. 243; Pfeiffer, Syn. Auric, no. 30; Mon. 
Auric, i. p. 36 ; Morch, Cat. Yoldi, p. 38 ; Mousson, Jouni. de 
Concli. 1869, p. 346 ; Martens & Langk. Don. Bism. p. .55 ; Gassies, 
Faun. Noiiv. Cal. p. 62 ; Pease, Jouru. de Conch. 1871, p. 93; 
Proc. Zool. Soc. 1871, p. 477 ; Paetel, Cat. Conch, p. 114 ; Schmeltz, 
Cat. Mus. Godeff. v. p. S8 ; Garrett, Proc. Phil. Acad. Nat. Sci. 
1879, p. 28; Journ. Phil. Acad. Nat. Sci. 1881, p. 402, 1885, 
p. 89. 

Easily distinguished by its large size (18 millim.) and uniform 
luteous colour. Abundant just above high-water mark ; it ranges 
from the Gambler Islands to the East Indies. 

2. Melampus fasciatus (Deshayes). 

Auricula fasciata, Deshayes, Encycl. Meth. ii. p. 90 ; Lam. Hist, 
viii. p. 337 ; Kiister, Auric, pi. .A., figs. 2, 3 ; Mousson, Java Moll, 
p. 46, pi. 5. figs. 28, 29. 

Melampus fasciatus, Beck, Ind. Moll. p. 107 ; (Tralia) H. & A. 
Adams, Proc. Zool. Soc. 1854, p. 1 1 ; Pt'eifFer, Syn. Auric, no. 33 ; 
Mon. Auric, i. p. 38; Mousson, Journ. de Conch. 1869, p. 348; 
Pease, Proc. Zool. Soc. 1871, p. 477; Martens & Langk. Don. 
Bism. p. 55; Paetel, Cat. Conch, p. 114; Schmeltz, Cat. Mus. 
Godeff. V. p. 88; Garrett, Journ. Phil. Acad. Nat. Sci. 1881, 
p. 402, 1885, p. 90. 

Conovulus fasciatus, Griffith, Cuv. Anim. Kingd. pi. 27. fig. 13 ; 
Anton, Verz. p. 48 ; Guerin, Icon. Moll. p. 17, pi. 7. fig. 8. 

Tralia (Pir a) fasciata, B.. & A. Adams, Gen. Moll. ii. p. 240. 

This, like the preceding species, lives just above high-watermark, 
and has the same extensive geographical range. 

It is subject to considerable variation in shape and colour. The 
type varies from bluish white to luteous, and is girdled with from 
four to six narrow chestnut bands on the body-whorl. Varieties of a 
uniform bluish-white, corneous, brownish, or orange-brown are not 
infrequent, as well as one of an orange-brown with three chestnut 
bands. The spire is marked with minute radiating grooves. 

3. Melampus parvulus, Nuttall. 

Melampus parvulus, Nuttall, MS., Pfeiffer, Syn. Auric, no. 11 ; 
Mon. Auric, p. 24 ; H. & A. Adams, Gen. Moll. ii. p. 243 ; Pease, 
Proc. Zool. Soc. 1871, p. 477 ; Martens & Langk. Don. Bism. 
p. 56, pi. 3. fig. 10; Paetel, Cat. Conch, p. 114 ; Brazier, Quart. 
Journ. Conch, i. p. 274. 

Common on the margins of mangrove-swamps. Mr. Brazier 
records it from Torres Straits. I took a few examples at Samoa and 
Wallis Islands. Mr. Nuttall obtained the type specimens at the 
Sandwich Islands. I have also received examples from New 
Caledonia. 

Proc. Zool. Soc— 1887, No. XX. 20 



286 MR. A. GARRETT ON THE TERRESTRIAL [.Mar. 1, 

The Viti shells, which are a little smaller than Sandwich-Island 
specimens, do not differ from the latter except in having in some 
examples one or two more denticles or plicae on the parietal wall, and 
the base more distinctly impressedly striated. It may be recognized 
by its ovate shaj)e, smooth shining surface, dark chestnut or olive- 
brown colour, short, convexly conoid spire, and mucronated apex. On 
the lower portion of the parietal region may be observed two approxi- 
mating folds, the lower one the smaller and occasionally wanting. 
There are usually one or two small denticles above, and the palate 
has five to seven laminae. The columclla-fold is continuous with 
the basal portion of the peristome. 

M. fjranum, Gassies, is either the same as M. purvulus or very 
closely related. 

4. Melampus tongaensis, Mousson. 

Melainpus tongaensis, Mousson, Journ. de Conch. 18/1, p. 22, 
pi. 3. fig. 8; Schmeltz, Cat. Mus. Godeff. v. p. 88; Pfeiffer, Mou. 
Pneum. (Auiic.) iv. p. 316. 

A number of examples were taken in the same station as the pre- 
ceding species. Dr. Graffe found the type specimens at Tongatabu, 
one ot the Tonga Islands. Prof. Mousson has described a variety 
j}a//i(/ula (/. c.) from Vavao in the same group. 

It is very closely allied to, and perhaps only a form of, M. parvulus. 
It is about the same size and colour, but is a little more oblong in 
shape and the si)ire more produced. The dentation and plicae are 
the same in the two species. 

5. IMelampus semisulcatus, Mousson. 

Melampus semisulcatus, Mousson, Journ. de Conch. 1869, p. 347, 
pi. 15. fig. 2; Paetel, Cat. Conch, p. 114; Schmeltz, Cat. Mus. 
GodefF. v. p. 88 ; PfeiflFtr, Mou. Pneum. (Auric.) iv. p. 318 ; Pease, 
Proc. Zool. Soc. 18/1, p. 477. 

Occurs on the n.argius of mangrove-swamps, where I gathered 
thousands of specimens. I also obtained it in similar stations at 
Upolu, one of the Samoa Islands. 

This species is of an oblong pear-shape and a uniform cinnamon 
colour; it has a short, usually eroded, spire, and is spirally grooved, 
the grooves being more or less evanescent on the middle of the body- 
whorl. There are three folds in the parietal region and usually two 
lamina in the palate. Length 11 niillim. 

6. Melampus sculptus, Pfeiffer. 

Melampus seulptus, Pfeiffer, Proc. Zool. See. 1859, p. 29; Men. 
Pneum. (Auric.) iv, p. 316. 

Melampus fricki, Pfeiffer, Proc. Zool. Soc. 1859, p. 29; Mon. 
Pneum. (Auric.) iv. p. 304 ; Pease, Proc. Zool. Soc. 1871, p. A77. 

Melampus semiplicafus. Pease, Proc. Zool. Soc. 1860, p. Hfi, 
1869, p. 60 (animal), 18/1, p. 477; Pfeiffer, Mon. Pneum. 
(Auric.) iv. p. 304; Schmeltz, Cat. Mus. Godeff. v. p. 88; Layard, 
Cat. Land and Freshw. Moll. N. Caled. p. 4. 



1887.] MOLLUSKS OF THE VITI ISLANDS. 287 

Melawpus strictus, Gassios, Journ. de Concli. 1874, p. 213 ; 
Pfeiffer, Mon. Piieutn. CAuric.) iv. p. 324. 

Melanipus pseudocommorlus, " Mousson," Schmeltz, Cat. Mus. 
Godeff. iv. p. 69 ; Paetel, Cat. Conch, p. 114. 

A few examples found just above high-water mark on a small 
islet on the north coast of Vanua Levu. 

Pfeiffer's tvpe specim.ens in Cuming's Museum were labelled 
Admiralty Islands. His M. fricki, together with Pease's semi- 
plicatus and INIousson's pseudncommodus, were obtained at the 
Sandwich Islands, where I first discovered Pease's type specimens. 
Shells received from New Caledonia labelled M, strictus do not 
differ from Viti examples. 

The longitudinal plications on the upper third of the shell, pale 
or dark brownish colour, numerous whorls, rather long acute spire, 
and single parietal fold will readily distinguish this species. The 
basal portion is also more or less distinctly plicated. The palate 
has from one to three laminae. Length 10 millim. 

7. Melampus consanguineus, sp. nov. 

Shell imperforate, solid, obovate, smooth, shining, faintly striated 
with lines of growth, light chestnut-brown ; spire couvexly conoid, 
apex mucronate ; sutuial line distinct, linear: whorls 7, flattened, 
the last one subangulate on the shoulder, and obliquely impressedly 
striated at the base ; the lower portion of the parietal region with 
two spiral plications, the upper one the larger, and occasionally 
there exists one or two posterior denticles; palate with from 11-14 
white plicae ; columellar fold continuous with the basal portion of 
the peristome. 

Length 9, diam. 5 millim. 

Not uncommon at high-water mark at Vanua Levu. 

The uniform pale chestnut colour and numerous plications in the 
throat will determine it. 

8. Melampus striatxjs. Pease. 

Melampus striatus (Tralia), Pease, Proc. Zool. Soc. 1861, p. 244, 
1871, p. 477; Pfeiffer, JVIon. Pneum. (Auric.) iv. p. 311 ; Garrett, 
Journ. Acad. Nat. Sci. Phil. 1885, p. 89. 

Melampus mordrouzieri, Souverbie, Journ. de Conch. 1866, 
p. 148, pi. 6. tigs. 1, la; Pfeiffer, Mon. Pneum. (Auric.) iv. 
p. 312. 

Melampus ornatus, Mousson, Journ. de Conch. 1871, p. 21, 
pi. 3. fig. 7 ; Pfeiffer, Mon. Pneum. (Auric.) iv. p. 312. 

Not infrequent on the margins of mangrove-swamps in the Viti, 
Tonga, and Samoa Islands. It also inhabits the Society Islands 
and New Caledonia. 

It may be distinguished by its oblong-ovate form, the brownish- 
corneous, chestnut-brown, or greenish-brown colour, its mucronated 
spire, and the 8 whorls marked by closely-set transverse impressed 
lines, the upper half with small longitudinal plications, which give 

20* 



28S MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

that part of the shell a granulated appearance. The transverse lines 
are frequently evanescent on the middle of the body-whorl. There 
are from two" to three folds on the parietal region, the upper one 
small and granuliform. There may he observed one to three 
lamelliforni plications in the palate, and sometimes several raised 
white parallel strise. Length 9-10 millim. 

M. yranifer, Mousson, an East-Indian species, is very closely 
allied to, if not identical with, M. striatus. 

9. Melampus adamsianus, Pfeiflfer. 

Melampus adamsianus, Pfeiffer, Proc. Zool. Soc. 1854, p. 121 ; 
Syn. Auric, no. 12; N^ovit. Conch, i. p. 18, y)l. 5. figs. 17-19; 
Mon. Auric, i. p. 24 ; Gassies, Faun. Nouv. Caled. p. 57, pi. 7. 
fig. 2 ; Hutton, Cat. Moll. New Zeal. p. 576 ("ex Pfeiffer ") 

Tralia adamsiana (Pira), H. & A. Adams, Gen. Moll. ii. p. 244. 

Melampus variabilis, Gassies, Faun. Nouv. Caled. p. 65, pi. 6. 
fig. 8 ; Pfeiffer, Mon. Pneum. (Auric-) iv. p. 315. 

Melampus cinereus, Gassies, Journ. de Conch. 1867, p. 62 ; 
Pfeiffer, Mon. Pneum. (Auric.) iv. p. 314. 

Melampus avenaceus, Mousson, Journ. de Conch. 1870, p. 134; 
1871, var. vavaoensis, p. 21. 

Melampus angustus, " Mousson," Schmeltz, Cat. Mus. Godefif. iv, 
p. 68; Paetel, Cat. Conch, p. 114 (juvenile). 

A small and very variable species, 7 to 10 millim. long, and of 
various colours — light or dark chestnut, luteous, fulvous, cinereous, 
frequently ornamented with bands and lines of a chestnut colour on 
a pale ground or pale bands on a dark ground. The shape varies 
from oblong-ovate to subcylindrical ; surface smooth, shining ; base 
with oblique incised strise. Spire short or elongate, acute, obscurely 
radiately plaited or grooved. Parietal region with from one to four 
pHcations, the posterior two when present granuliform. Columellar 
fold more or less continuous with the peristome. The variety 
vavaoensis is common in the Viti group, associated with the type, 
into which it gradually intergrades. 

Specimens occur in great profusion just above high-water mark in 
sheltered places. Dr. Griiffe found it in the Tonga group and it 
is abundant in New Caledonia, whence I have received numerous 
examples labelled M. adamsianus, M. cinereus, and M. variabilis. 
The New-Caledonian shells exhibit the same variation as the Viti 
shells, some of which have the spire so much elongated that they 
might easily be mistaken for a distinct species ; but having carefully 
studied several thousand specimens collected in the latter group, I 
find the character individual only. The number of plications in the 
aperture cannot, except in certain species, be relied on as a specific 
character. It was first described from specimens in the Cumingian 
Museum, and the habitat "New Zealand" is probably erroneous. 
Mr. Hutton, in his Catalogue of New-Zealand MoUusca, merely 
repeats Pfeiffer's description. Von Martens does not include it in 
his list of New-Zealand shells. 



1887.] MOLLUSKS OF THE VITI ISLANDS. 289 

10. Melampus crebristriatus, sp. nov. 

Shell imperforate, solid, obovate, slightly shining, striated with 
lines of growth and marked by rather crowded transverse incised 
lines ; colour dark chestnut-brown or fulvous, with or without two 
light chestnut bands ; spire mucronate, convexly conoid ; suture 
distinct, linear ; whorls 7—8, subplanulate, the last one subangulate 
on the shoulder ; tbe lower portion of the parietal region with two 
subcoutiguous folds, the lower one small, and occasionally there 
exists small posterior denticles; palate with from 10-18 whitish 
laminae on a layer of whitish callus ; peristome and columella 
fulvous. 

Length 10-13, diam. 6-7 millim. 

A few examples found near high-water mark on the north coast 
of Vanua Levu. 

11. Melampus rusticus, sp. nov. 

Shell small, imperforate, obovate, finely striated, brown, with 
irregular longitudinal fulvous stripes and small spots ; spire short, 
conoid, apex eroded, truncate ; whorls 4 remaining, last one sub- 
angulated, obliquely striated at the base ; aperture elongate, narrow, 
slightly oblique, violaceous or brown, base rounded ; parietal region 
with an acute horizontal lamina just below the middle ; columellar 
fold sharp, oblique, and continuous with the acute peristome ; palate 
with 4-6 faint laminae. 

Length 7, diam. 5 millim. 

Rather common on the margins of mangrove-swamps. I also 
found it in the Tonga and Samoa Islands. 

It is the same shape as, but smaller than, M. semisulcatus, with 
which it is found associated. It is also darker coloured, and differs 
from the latter in the absence of spiral sulcations and in having only 
one parietal fold. 

12. Melampus incisus, sp. nov. 

Shell imperforate, obconic, solid, marked by fine incremental 
striae and spiral incised lines, which are sometimes evanescent on 
the middle of the body-whorl ; colour brown or luteous, with or 
without four transverse chestnut bands, and frequently with irregular 
longitudinal more or less interrupted fulvous lines and dots ; spire 
short, conoid, apex eroded, truncate ; whorls 5 remaining, last one 
subangulated above ; suture linearly impressed and slightly lacerated ; 
aperture somewhat oblique, elongate, violaceous brown ; parietal 
region with two contiguous folds just above the columellar plait, the 
upper one the larger, above which are from two to six more or 
less distinct denticles ; palate with five to fourteen white irregular 
laminae ; columellar fold oblique, continuous with the peristome. 

Length 8-10 millim. 

Not infrequent on the margins of mangrove-swamps in Vanua 
Levu. 



290 MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

Genus Tralia, Gray. 

1. Tralia melanostoma (Garrett). 

Persa melanostoma, Garrett, Amer. Journ. Conch. 1872, p. 224, 
pi. 19. fig. 11 ; Schmeltz, Cat. Mus. Godeff. v. p. 87. 

Melampus melanostoma, Pfeiffer, Mod. Pneum. (Auric.) iv. 
p. 325. 

Abundant and gregarious under stones, near and a little below 
high-water mark, ou the east end of Taviuni Island. 

A small oblong-ovate or elliptically ovate tawny-brown species, 
with a blackish aperture, short, acute, spirally striated spire, and 
generally with a transverse brown band beneath the suture. Aperture 
rounded at the base, narrow above. Parietal region with one or two 
superior denticles, and a large fold just above the columellar plait. 
Peristome thick, labiated within and sinuous above. Length 4| 
millim. 

2. Tralia costata (Qnoy & Gaimard). 

Auricula costata, Qiioy & Gaimard, Voy. Astrol. ii. p. 173, pi. 13. 
figs. 43-46 ; Deshayes, Lam. Ilist. viii. p. 337 ; Kiister, Auric, 
p. 46, pi. 7. figs. 5-7. 

Melampus cosiatus, Beck, Ind. Moll. p. 107 ; {Tralia) H. & A. 
Adams, Proc. Zool. Soc. 1854, p. 12; Pfeiffer, Syn. Auric, no. 56 ; 
Mon. Auric, i. p. 55 ; Mousson, Journ. de Conch. 1870, p. 135 ; 
Paetel, Cat. Conch, p. 114. 

Tralia costata {Persa), H. & A. Adams, Gen. Moll. ii. p. 245 ; 
Chenu, Man. Conch, i. p. A77. fig. 352/. 

Common, associated with the preceding species. 

A solid, ovate, longitudinally ribbed, fulvous or reddish-brown 
species, with three plaits on the parietal wall and columella. The 
peristome is thick and sinuous above. Length 8-10 millim. 

3. Tralia alba (Gassies). 

Melampus alius, Gassies, Journ. de Conch. 1865, p. 21 1 ; Pfeiffer, 
Mon. Pneuin. (Auric.) iv. p. 326. 

Melampus luiithis. Pease, Amer. Journ. Conch. 18G9, p. 75 ; 
Pfeiffer, Mon. Pneum. (Auric.) iv. p. 327. 

Melampus pellucidus, Pease, Journ. de Conch. 1871, p. 93. 

Two specimens occurred to my notice under a clump of coral on 
the east coast of Taviuni Island. 

A smootii white or horn-culoured species, 3g to 5 millim. long, of 
an oblong-ovate form, with a rather j)roduced spire, and the plica- 
tions the same as in the preceding species. 

Genus Laimodonta, Nuttall. 

I. Laimodonta layardi, H. & A. Adams. 

Ophicardelus layardi {Laimodonta), II. & A. Adams, Proc. Zool. 
Soc. 1854, p. 35. 



188/.] MOLLTJSKS OF THE VITI ISLANDS. 291 

Laimodonta layardi, H. & A. Adams, Gen. Moll. ii. p. 246. 

Melampus layardi, Pfeiffer, Syn. Auric, no. 48 ; Mon. Auric, i. 
p. 51 ; Gassies, Faune Nouv. Caled. p. 61, pi. 7. fig. 7; Tenneiit's 
Ceylon, i. p. 239; Cox, "Exchange List," p. .33; H. Nevill, Eiium. 
Hei. etc. Ceylon, 1871, p, 4. 

Laimodonta conica. Pease, Prnc. Zool. Soc. 1862, p. 242 ; Amer. 
Journ. Conch. 1868, p. 101, pi. 12. fig. 15 ; Journ. de Concli. 
1871, pp. 93, 94 ; Proc. Zool. Soc. 1871, pp. 470, 477 ; Schmeltz, 
Cat. Mus. Godeff. v. p. 81 ; Garrett, Journ. Phil. Acad. Nat. Sci. 
1881, p. 403, 1885, p. 91. 

Lnemodonta conica. Martens & Langk. Don. Bism. p. 57, ph 3. 
fig. 13. 

Laimodonta anaaensis, Mousson, Journ. de Conch. 1869, p. 63, 
pi. 5. fig. 1. 

Plecotrema anaaensis, Paetel, Cat. Couch, p. 114. 

Melampus conicus, Pf'eitfer, Mon. Pneum. (Auric.) iv. p. 319. 

"? " Melampus anaaensis, Pieiffer, I. c. p. 320. 

A few dead examples found in beach-sand. 

Since the publication of my paper on the Land-shells of Cook's 
Islands Mr. E. L. Layard has sent me for determination a Ceylon 
species of Laimodonta, which proves to be L. layardi, and is 
identical with Pease's L. conica. Dr. Cox and Gassies record 
L. layardi from New Caledonia, and Pease quotes it {conica) from 
" Central Pacific." I have obtained it in all the groups from the 
Paumotus to the Viti Isles. 

The species now under consideration is closely allied to the 
Sandwich-Island L. bronni, but is smaller, more slender, and the 
spiral engraved lines are more conspicuous. My examples average 
from 6 to 8^ millim. in length. Colour chestnut-brown, with one or 
two whitish bands. Tiie outer lip is slightly sinuous posteriorly 
and has one or two internal riblets. All the three descriptions 
alluded to mention a single plait in the palate. In the eight 
specimens before me six have two ril)lets in the palate. 

Station under stones above high-water mark. 

Genus Pedipes, Adanson. 
1, Pedipes jouani, Montrouzier. 

Pedipes jouani, "Montrouzier," Souverbie, Journ. de Conch. 
1862, p. 244, pi. 9. fig. 1 1 ; Gas>ies, Fauu. Nouv. CaleJ. p. 6.5, pi. 6. 
fig. 22; Pfeiffer, Moa. Pneum. (Auiic.) iv. p. 332. 

Pedipes subylobosus, Garrett, Proc. Phil. Acad. Nat. Sci. 1873, 
p. 236, pi. 3. tig. 70. 

Eight examples found lurking under stones a little below high- 
water mark at Laiithala Island. 

Our specimens do not differ from the New-Caledonian shells except 
in being |)aler. Its subgbibose form, small, crowded granulated 
spiral ridges, fulvous-brown colour, slightly shouldered boilv-whorl, 
and short miicrouated spire will readily iHstinguish it. The flattened 
callose columella is armed with two compressed transverse folds. 



292 MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

above which may be observed a sharp deflected jiarietal plait, 
and a tubercle on the inner margin of the acute peristome, which 
latter is thickened within. Length 4| millim. 

Genus Pythia, Bolten. 

1 . PvTHiA POLLEX (Hinds). 

Scarains 2)oHex, Hinds, Ann. Nat. Hist. \. p. 82 ; Voy. Sulph., 
Zool. p. (50, pi. 16. figs. 9 & 10 ; A. Adams, Proc. Zool. Soc. 18.50, 
p. 1.50 ; Ann. Nat. Hist. 2nd ser. viii. p. G9 ; Reeve, Conch. Icon, 
sp. 7, fig. 7. 

Fythia pollex, Pfeiffer, Syn. Auric, no. 82 ; Mon. Auric, i. p. 86 ; 
Brit. Mus. Cat. Auric, p." 65 ; H. & A. Adams, Gen. Moll. ii. 
p. 240; Mousson, Joiirn. de Conch. 1870, p. 133; Paetel, Cat. 
Conch, p. 114 ; Schmeltz, Cat. Mus. Godeff. v. p. 87; Cox, Proc. 
Linn. Soc. New South Wales, vi. p. 611. 

Scarabus zonatus, Hombron & Jacquinot, Voy. Pole Sud, Zool. 
V. p. 41, pi. 10. figs. 18-20. 

Very abundant and generally distributed throughout the group. 
Occurs beneath decaying vegetation in forests near the sea-shore. 

This species is subject to considerable variation in size, shape, 
and colour. Though usually unibilicated, it is nevertheless very 
frequently imperforate. The shape varies from broad ovate to 
oblong ovate ; spire subacute, more or less produced, and laterally 
subangulated. The sculpture consists of longitudinal, closely set, 
elevated striae, often evanescent on the body-whorl, and very con- 
spicuous and slightly arched on sjiire and upper part of the last 
•whorl. The superior parietal tooth is subtriangular, the lower one 
compressed, fold-like, and subdu[)licated. The columellar jilait is 
slightly oblicpie, compressed, and in imperforated specimens is con- 
tinuous with the broadly expanded and shghtly reflected peristome. 
The palate is armed with two stout and from four to six small 
teeth. 

The colour varies from light chestnut to blackish chestnut, more 
or less consjiicuously mottled with luteous, and generally with one 
or two pale transverse bands above, The varices, which are not very 
conspicuous, are usually spotted with white or luteous. Uniform 
horn-coloured or luteous specimens with or without chestnut 
mottlings are not uncommon. Aperture white or buff-yellow, with 
or without chestnut maculations. Sometimes the very dark examples 
show three or four pale transverse bands. The following measure- 
ments will show the variation in shape and size: — 

Length 36, diam. 21 millim. 
>> "^4, „ Zo ,, 

j» 23, ,, 15 ,, 

2. Pythia albovaricosa, Pfeiffer. 

Pythia aUovaricosc, Pfeiffer, Zeit. Malak. 1653, p. 190; Syn. 
Auric, no. 84 ; Mon. Auric, i. p. 87 ; Brit. Mus. Cat. Auric, p. 66 ; 



1887.] MOLLUSKS OF THE VITI ISLANDS. 293 

Novit. Conch, i. p. 6, pi. 3. figs. 1 & 2 ; H. & A. Adams, Gen. Moll. 
ii. p. 240 ; Cox, Proc. Linn. Soc. New South Wales, vi. p. 592. 

Scarabus albovaricosus. Reeve, Conch. Icon. sp. 4, pi. 1. figs. 
2, 6. 

In looking over a lot of about 200 specimens of P. pollex, I found 
amongst them an example of Pfeiflfer's P. albovaricosa, which does 
not differ in a single feature from five Solomon-Island s])ecimens 
received from Dr. Cox. I cannot indicate the island whence tiie 
shell was obtained, but am inclined to believe it was Kantavu. 

Dr. Pfeiffer and Reeve, on the authority of Cuming, cite Celebes 
as habitat of this species ; and Dr. Cox, in his valuable paper on the 
" Nomenclature and the Distribution of the genus Pi/t/iia," says it 
is a common Solomon-Island species. As it is now well ascertained 
that many of Cuming's localities are erroneous, the former habitat 
needs confirmation. 

This species may be characterized by its large size (31 to 44 
millim.), rather light texture, oblong-ovate form, smooth bodv- 
whorl, the upper part of which, together with the spire, is marked 
by short, longitudinal, slightly arcuated grooves. Colour light or 
dark chestnut, sometimes light fulvous with very small darker 
irrorations. Tlie varices are white with wide black or dark chestnut 
margins, and the six specimens now before me all have a large 
lateral diffused blackish patch on the front and back of the body- 
whorl. Aperture luteous or whitish. Palatal teeth 4 or 5. 

3. Pythia savaiensis, Mousson. 

Pythia pantherina, A. Adams, var. uveana, Mousson, Journ. de 
Conch. 186.5, p. 1/7 ; Sclimeltz, Cat. Mus. Godeff. iii. p. 2S ; Pease, 
Proc. Zool. Soc. 1871, p. 477; Paetel, Cat. Conch, p. 114 ; PfeifTer, 
Mon. Pneum. (Auric.) iv. p. 348 ; Cox, Proc. Linn. Soc. New 
South Wales, vi. p. 617. 

Pythia savaiensis, Mousson, Journ. de Conch. 1869, p. 345, 
1870, p. 133; Pease, Proc. Zool. Soc. 1871, p. 477; Schmeltz, 
Cat. Mus. Godeff. v. p. 87 ; Pfeiffer, Mon. Pneum. (Auric.) iv. 
p. 341 ; Cox, Proc. Linn. Soc. New South Wales, vi. p. 613; G. 
Nevill, Hand-list Moll. Mus. Calcutta, p. 223. 

Not uncommon in forests near the sea-shore at Kioa Island. Dr. 
Griiffe obtained it at Ovalau, Samoa, and Tonga group. I gathered 
several hundred examples at Wallis Island (=Uvea or Uea), one of 
the latter group. 

A solid, ovate, umbilicated (rarely imperforate) species, 21 to 28 
millim. long, with the striation of P. pollex, and the lower parietal 
fold simple ; the n])per one is small and trilobed. Colour corneous 
or yellowish-horn colour mottled with chestnut, rarely uniform light 
or dark chestnut. Varices spotted with white, and the aperture is 
luteous. The palate is furnished with 4 or 5 teeth. 

The small size, absence of bands, uniform shape, and the simple 
lower ]»arietal fold will readily separate it from P. pollex. 

Mousson's name savaiensis is derived from Savaii, one of the Samoa 



294 MR. A. GARRETT ON THE TERRESTRIAL, [Mar. 1, 

Islands. He very correctly drops one i ; and Pease, who retains both, 
spells it " savaiiensis." Both Schmeltz and Paetel erroneously 
quote it as " savai/ensis." All, however, refer to one and the same 
species. 

4. PvTHiA LENTiGiNosA, Garrett. 

Pijthia lentiyinosa, Garrett, Amer. Journ. Couch. 18/2, p. 220, 
pi. 19. fig. 4; Schmeltz, Cat. Mus. Godeff. v. p. 87; Pt'eiffer, Mou. 
Pneum. (Auric.) iv. p. 337 ; G. Nevill, Hand-Hst Mus. Calcutta, 
p. 222 ; Cox, Proc. Linn. Soc. New South Wales, vi. p. 604. 

This species occurred to me in only one location, a small islet close 
to the east end of Taviuni, where it was found in abundance associated 
with P. pollex. 

It is remarkably uniform in colour, and differs but little in shape 
and size. It is a solid, ovate or oblong-ovate shell, 21 to 29 millim. 
long, yellowish white, profusely spotted with small fulvous-brown 
maculations, and with a honey-yellow aperture. Tiie varices, which 
are not very conspicuous, are spotted with wiiite and chestnut, which 
on the sides of the body-whorl are elongated into stripes. The basal 
perforation varies from rimate to umbilicate. The lower parietal 
fold shows a very slight indication of an external groove. 

5. Pythia perovata, Garrett. 

Pythia perovata, Garrett, Amer. Journ. Conch. 1872, p. 221, 
pi. 19. fig. 5 ; Schmeltz, Cat. Mus. Godeff. v. p. 87 ; Pfeiffer, Mon. 
Pneum. (Auric.) iv. p. 34 ; Cox, Proc. Linn. Soc, New South Wales, 
vi. p. 608. 

I collected about 2000 specimens of this species on the margins of 
mangrove-swamps at Vuna Dawa on the north side of Natawa Bay, 
Vanua Levu, and at Na Viti Levu Bay in the N.E. part of Viti 
Levu. Examples taken at the latter location were larger than the 
Vuna Dawa siiells. 

Tb.is ver}' distinct species is of an ovate or abbreviate-ovate shape, 
solid ; spire short, base imperforate, sometimes rimate, more rarely 
perforated ; longitudinally striated, the striae straight and most 
conspicuous above. The dentation is white or tawny ; the upper 
tooth on the parietal wall is vertical, elongate, crest-iike, simple or 
very rarely with a slight inferior lobe j)ri>jecting to the lelt. The 
lower fold is compressed and fiirni>hed with a small, short, tooth-like 
duplication. The columelhir plait is obliquely twisted and continuous 
with the widely expanded peristome, which latter is simple above 
and sli;:htly reflected below. Palate with four, rarely five teeth, two 
of which are t!ie larger. Colour light to dark chestnut or reddish 
chestnut, rarely light yellowish-horn colour, frequently indistinctly 
mottled with a tint daiker than the ground-colour, and very olten 
with a blackish sutural band. Varices rarily spotted with whitish. 
Lengtli 15 to 24 millim. The adults are very frequently eroded 
over the whole surface. 



1887.] MOLLUSKS OF THE VITI ISLANDS. 295 

Genus Plecotrema, H. & A. Adams. 

1. Plecotrema souverbiei, Montrouzler. 

Plecotrema souverbiei, " Montrouzier," Souverbie, Journ. de 
Conch. 1S62, p. 24G, pi. 9. fig. 12 ; Gassies, Faune Nouv. Cale'd. 
p. 67, pi. 6. fig. 23 ; Pt'eiffer, Mon. Pueum. (Auric.) iv. p. 343. 

Plecotrema turrita, Garrett, Proc. Phil. Acad. Nat. Sci. 1873, 
p. 235, pi. 3. fig. 68. 

Not uncommon under stones a little below high-water mark on 
the east end of Taviuni. It also inhabits New Caledonia. 

This remarkable species is easily recognized by its ovate-conical 
form, scalariform spire, corneous or dull fulvous colour, subper- 
forated and angulated base. The sculpture consists of numerous, 
small, crowded, granulated spiral ridges, with the intermediate 
grooves crossed by sublaminated striifi. There is a prominent crest- 
like varixjust behind the peristome, which latter is acute, continuous, 
and slightly porrected. Parietal region with a superior nodiform 
tooth beneath, of which there is a simple acute plait. Columellar 
fold small and nearly transverse. Outer lip labiate within aud 
bidentate. Length 4| millim.. 

Very closely allied to, if not identical with, P. bella, a Philippine 
species. 

2. Plecotrema hirsuta, Garrett. 

Plecotrema hirsuta, Garrett, Amer. Journ, Conch. 1872, p. 219, 
pi. 19. fig. 2 ; Schmeltz, Cat. Mus. Godeff. v. p. 87 ; Pfeiffer, Mon. 
Pueum. (Auric.) iv. p. 348. 

Abundant in the same station and locality as the preceding 
species. 

An imperforated, solid, acutely-ovate, corneous species, with spiral 
rugose lirse, 16 to 18 on the body-whorl, garnished with short, 
deciduous, curved hair-like setse. The intervening sulcations deep, 
narrower than the lirations, and crossed by sublaminated striae. 
There is a stout obtuse varix just behind the peristome, and the 
dentation, excepting the lower parietal fold, which is bifid, does not 
differ from the preceding species. Length 5 to 7 millim. 

3. Plecotrema octanfracta (Jonas). 

Pedipes octanfracta, Jonas, Zeit. Malak. 1845, p. 160. 

Plecotrema octanfracta, Jerkeli, Nachr. Malak. Ges. 1872, p. 65 ; 
Pfeiffer, Mon. Pneum. (Auric.) iv. p. 34G. 

Plecotrema cluusa, H. & A. Adams, Proc. Zool. Soc. 1853, p. 121 ; 
Gen. iMoU. ii. p. 241. ; Pfeiffer, Syn. Auric, no. 101 ; Novit. Conch. 
i. p. 15, pi. 5. figs. 9-11 ; Mon. Auric, i. p. 103 ; Pease, Proc. Zool. 
Soc. 1871, pp. 4;"j9, 477; Paetel, Cat. Conch, p. 144; Schmeltz, 
Cat. Mus. Godeff. v. p 87. 

Plecotrema coasobrina, Garrett, Proc. Phil. Acad. Nat. Sci. 1873, 
p. 236, pi. 3. fig. 69. 

Not uncommon under stones a little below high-water mark at Kioa 



29C MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

and Taviuni. It also iiiliabits the Sandwich and Paumotu Islands, 
and I found it very abundant and gregarious at the Gambler Islands. 
A small, solid, ovate species with the spiral hra; of P. liirsuta, 
but more numerous, smoother, more crowded, and the aperture is 
not so much contracted. The external varix is smaller, and the 
base of the shell is more rounded than in the latter sjjecies. Colour 
light brownish, sometimes corneous, usually with a faint jjale zone 
beneath the suture, and the aperture is more or less tinged with 
brownish. The dentation is the same in both species. Length 3 
to 5 millim. 

Genus Cassidula, Forussac. 

1. Cassidula intuscarinata, Mousson. 

Auricula {Cassidula) intuscarinata, Mousson, Journ. de Conch. 

1870, p. 132, pi. 7. fig. 9. 

Cassidula intuscarinata, Schmeltz, Cat. Mus. Godeff. v. p. 88 ; 
Pfeiffer, IMon. Pneuni. (x\uric.) iv. p. 3.53. 

Occurs in profusion on the mud in mangrove-swamps at Viti 
Levu and Vanua Levu. I have received the same sjjecies from 
New Caledonia labelled C. mvstellina. 

This species, like nearly all the shells inhabiting swamps, is very 
frequently stained and more or less eroded. "When in good condition 
it is of a brown or brownish-liver colour, sometimes olive-brown, with 
an obscure pale band on the subangulated shoulder. Karely light 
fulvous-brown, with an indistinct light chestnut band between the 
shoulder and the sutural line. The base, aperture, and the peristome 
tawny flesh-colour, the latter with a stout external varix. The 
shape of the bhell is subovate, \\'\X.\\ s])iial incised lines, and the 
apertiu-e is obauriform. The strongly labiated hp is deeply emargi- 
nated above. Upper parietal tooth small, nodiform, and the plait 
beneath is neaily transverse. Coluniellar fold slightly oblique. 
Length 14 to 20 miUim. 

2. Cassidula crassiuscula, Mousson. 

Cassidula crassiuscula, Mousson, Journ. de Conch. 1869, p. 343, 
pi. 15. fig. 1 ; Paetel, Cat. Conch, p. 114; Pease, Proc. Zool. See. 

1871, p. 477 ; Schmeltz, Cat. Mus. Godeff. v. p. 88 ; Pfeiffer, Mou. 
Pnenm. (A.uric.) iv. p. 352. 

Auricula {Cassidula) crassiuscula, Mousson, Journ. de Conch. 
1871, I>. 191. 

Cassidula 7iucleus, G&ssies (Martvn ?), Faune Nouv. Caled. p. 71, 
pi. 3. fig. 9. 

Like the preceding species, it occurs in profusion on the mud in 
mangrove-swamps, but is more generally diffused throughout the 
group. It also iuhabits Tonga and the Samoa Islands. Mr. Layard, 
of New Caledonia, sent me specimens collected in that island, 
labelled " Melampus nucleus, Martyn." 

It has the same shape and dentation as C. intuscarinatus, but is 
much more variable in colour, and in size ranges from 10 to Id 



1887.] MOLLUSKS OF THE VITI ISLANDS. 297 

millim. long. It is of different shades of chestnut-brown, white, 
corneous, fulvous, frequently witli from one to four pale transverse 
bands on the body-whorl, and more rarely with a sutural livid band. 
Aperture pale fulvous, brownish or white, and the lip light fulvous 
or white. 

I am inclined to believe this species is diffused throughout the 
New Hebrides, Solomon Islands, and, perhaps, extends its range into 
the East Indies. I am sure Samoa is the eastern limit of the genus 
Cassidula ; Martyn's C. nucleus, which is erroneously quoted as a 
Tahitian species, does not inhabit that group. 

1 am also strongly inclined to believe Mousson's O. crassiuscula 
is identical with his C. sulculosa, an East-Indian species. 

3. Cassidula paludosa, Garrett. 

Ophicardulus paludosus, Garrett, Amer, Journ. Conch. 1872 
p. 220, pi. 19. fig. 3. 

Cassidula paludosa, Paetel, Cat. Conch, p. 114. 

Plecotrema paludosa, Schmeltz, Cat. Mas. Godeff. v. p. 87. 

Melampus paludosus, Pfeiffer, Mon. Pneum. (Auric.) iv. p. 327. 

Common in the mud in mangrove-swamps, and probablv generally 
distributed in the group. I took a \'q\\ examples at Up'olii, one of 
the Samoa Islands. 

A small, solid, ovate, whitish, yellow-corneous, or chestnut-brown 
species, with fine, spiral, incised, punctured lines and a carinated 
base. Spire convexly-conical, and more produced than in the 
preceding species. Aperture white, tawny, or fulvous, with the 
dentation of O. crassiuscula, with the addition, in old specimens, of a 
small denticle in the labial sinus. Length 8 to 10 millim. 



Genus Auricula, Lamarck. 

1. Auricula subula, Quoy & Gaimard. 

Auricula subula, Quoy & Gaimard, Voy. Astrol. ii. p. 171, pi. 13. 
figs. 39 & 40 ; Deshayes, Lam. Hist. viii. p. 334 .; Kuster, Auric, 
p. .5.3, pi. 8. figs. 9 & 10 ; Jay, Cat. Shells, 1850, p. 265 ; A. Adams 
& Reeve, Voy. Samarang, p. 55, pi. 14. fig. 15, with animal ; Pfeiffer, 
Auric, no. 147; Mon. Auric, i. p. 141; Gassies, Faune Nouv, 
Caled. p. 69 ; Paetel, Cat. Conch, p. 115. 

Pythia subula. Beck, Ind. Moll. p. 104. 

Ellobium subula, H. & A. Adams, Proc. Zool. Soc. 1853, p. 8 • 
Gen. Moll. ii. p. 238, pi. 82. fig. 1. 

Auricula elongata, " Parreyss," Kuster, Auric, p. 53, pi. 8. 
figs. 6-8; Jay, Cat. Shells, 1850, p. 264 ; Pfeiffer, Auric, no. 146; 
Mon. Auric, i. p. 140; Morelet, Ser. Conch, p. 93; Mousson, 
Journ. de Conch. 1871, p. 18 ; Paetel, Cat. Conch, p. 1 15 ; Schmeltz, 
Cat. Mus. Godeff. v. p. 88. 

Auricula buddii, Parreyss, MS. 

Ullobium elonyatum, H. & A. Adams, Proc. Zool. Soc. 1854, p. 8 ; 
Gen. Moll. ii. p. 237. 



298 MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

Ellohium oparicum, H. & A. Adams, Proc. Zool. Soc. 18.54, p. 9 ; 
Gen. Moll. ii. p. 237. 

Auricula oparica, Pfeiffer, Syn. Auric, no. 46 ; Novit. Conch, i. 
p. 28, pi. 7. figs. 14-lG ; Mon. Auric, i. p. 139. 

AuricuJus subu/a, Pt'eilfer, Mon. Pneum. (Auric.) iv. p. 360. 

Auriculus clongatiis, Pfeiffer, I. c. 

Ain'iculus oparicus, Pfeiffer, /. c. 

A very aliuiidant species, inhabiting the margins of mangrove- 
swamps, and widely diffused throughout the group. Likewise 
common to the Tonga and Samoa Islands, and generally distributed 
over Melanesia. It has also been found at different points in the 
East Indies; and Morelet records it from Mauritius. 

A small specits, 9 to 16 miliim.long, of a slender fusiform shape ; 
smooth, shining, longitudinally striated, acute, with a convexly- 
conical spire, very frequently truncated by erosion, and more or 
less lacerated at the suture. Body-whorl narrow, usually longer 
than the spire, attenuated or rounded at the base, rarely rimate. 
Aperture elongate, white, or light fulvous, sometimes livid, with a 
compressed subtransverse plait on the lower part of the parietal wall, 
and two small, oljlique, approximating folds on the columella, the 
upper one sometimes evanescent. Peristome obtuse, in old specimens 
slightly sinuous above, and adnate next the suture. Colour white, 
beneath an epidermis which varies from pale olivaceous horn-colour 
to chestnut- black. 

A careful comparison of the descriptions of A. elongata and 
A. oparica has convinced me that they do not differ from y/. subula, 
which Quoy obtained at the New Hebrides. Pfeiffer, in his descrip- 
tion of A. elongata, mentions only a single columellar fold, and quotes 
the Sandwich Islands, "Feejee," and one of the Philippines as 
habitat. It certainly does not live on the former group. Schmeltz 
cites one of the Caroline Islands and "Tahiti," the latter erro- 
neous. A. oparica, which was described from specimens in Cuming's 
Museum, is assigned to " insula Opara (ins. Societatis)." There is 
no island of that name in the Society group ; but there is a very 
small island, about 600 miles south of Tahiti, called Rapa-Oparee, 
which from its small size and rugged surface is not likely to be the 
home of the marsh-loving Auriculce. 

2. Auricula semisculpta, H. & A. Adams. 

Ellohium semisculptum, H. & A. Adams, Proc. Zool. Soc. 1854, 
p. 9 ; Gen. Moll. ii. p. 237. 

Auricula semisculpta, Pfeiffer, Syn. Auric, no. 139 ; Mon. Auric, 
i. p. 136 ; Novit. Conch, i. p. 39, pi. 10. figs. 7-9 ; Gassies, Faune 
Nouv. Caled. p. 70, pi. 3. fig. 11; Schmeltz, Cat. Mus. Godeft'. v. 
p. 88. 

Auriculus semisculptus, Pfeiffer, Mon. Pneum. (Auric.) iv. 
p. 359. 

I found several hundred examples of this species in different 
parts of the group. They were all found buried in rotten bogs on 



1887.] MOLLUSKS ON THE VITI ISLANDS. 2U9 

the margins of mangrove-swamps. I also took many in similar 
stations at Wallis Island and Samoa. I have received it from New 
Caledonia, and, undoubtedly, it ranges throughout Melanesia. The 
locality "Gambler Islands," usually assigned to this species, is 
decidedly erroneous. There are no swampy lands and not a single 
perennial stream in the group. Schmeltz is also wrong in citing 
Huaiiine, Society Islands, as habitat. 

This species varies considerably in shape, thickness, and in size 
ranges from 12 to 30 millini. long. The surface is shining, longi- 
tudinally striated, and the upper portions of the whorls are 
sculptured by crowded spiral rows of minute granules, which, in 
large adults, sometimes cover the whole surface of the body-whorl. 
The lower part of the parietal region is aimed with a prominent, 
compressed, oblique fold, and just beneath is a smaller and more 
vertical one on the columella. The peristome is rather strongly 
labiated and sinuous above. The shape of the shell varies from 
oblong-ovate to oblong-tnrreted, the spire is more or less produced, 
and the base impeiforated. Colour white, beneath a fulvous-yellow 
or yellowish horu-coloured epidermis. 

Genus Truncatella, Risso. 
1. Truncatella valida, Pfeiffer. 

Truncatella valida, Pfeiffer, Zeit. Malak. 1846, p. 182; Mon. 
Auric. (Appendix) i. p. 184 ; Jav, Cat. Shells, 18.54, p. 253 ; Kiister, 
Mon. p. 11, pi. 2. figs. 7, 8, 19-21, 23; H. & A. Adams, Gen. 
Moll. ii. p. 311 ; Martens, Ostas. Zool. ii. p. 262; Paetel, Cat. 
Conch, p. 118 ; Pease, Proc. Zool. Soc. 1871, p. 477. 

Truncatella vitiana, Gould, Proc. Bost. Soc. Nat. Hist. 1847, 
p. 208; Expl. Exp., Shells, p. 109, fig. 126; Otia Conch, p. 40; 
H. & A. Adams, Gen, Moll. ii. p. 311 ; Pfeiffer, Mon. Pneum. ii. 
p. 6; Mousson, Journ. de Conch. 1869, p. 356, 1870, p. 195; 
Paetel, Cat. Conch, p. 118; Schmeltz, Cat. Mus. Godeff. v. p. 104. 

Truncatella vitiacea, Mousson, Journ. de Conch. 1865, p. 185. 

Taheitia vitiana, Pease, Proc. Zool. Soc. 1871, p. '^77' 

Truncatella conspicua, " Bronn," Pfeiffer, Mon. Auric. (Appendix) 
i. p. 184 ; Paetel, Cat. Conch, p. 118; Layard, Cat. Land & Fresh- 
water Shells N. Caled. p. 1. 

This species occurs in abundance in all parts of the group, and 
lives just above high-water mark in sheltered places. It occurs also 
in the Samoa, Tonga, and Ellis group, and is diffused throughout 
Australasia and the East-India Islands. 

It may be characterized by its solid texture, slightly tapering 
cylindrical form, white, luteous, corneous, or ruddy corneous colour, 
and 4| slightly convex persistent whorls. The sculpture consists of 
nearly erect, obtuse ribs (25 to 35) on the body-whorl, and the base 
is more or less distinctly carinated. The peristome is thick, slightly 
expanded and auriculated at the suture. The operculum is thin, 
convex, smooth, with an elastic lamina-like margin. 

Length 6 to 8 millim. 



300 MR. A. GARRETT ON THE TERRESTRIAl, [Mar. 1, 

I am inclined to believe that T. pacifica. Pease, from the Caroline 
Islands, is identical with T. valida. 

2. Truncatella rustica, Mousson. 

Tnmcatella rustica, Mousson, Journ. de Conch. 1865, pi. 14. 
fig. 8; Paetel, Cat. Conch, p. 118; Schmeltz, Cat. Mus. Godeff. v. 
p. 104 ; Pfeiffer, Mon. Pneum. iv. p. 14. 

Truncatella costellifera. Pease, Proc. Zool. Soc. 1871, pp. 468, 
A77 ; PfeiflFer, Mon. Pneum. (Auric.) iv. p. 16. 

I found a few examples of this species at Taviuni, associated with 
T. vitiana. It was also obtained at "Wallis Island ( = " Uvea"), 
where Dr. Griiffe found the type specimens. Mr. Pease's T. costel- 
lifera, which Mr. Brazier obtained at Vavau, Tonga Islands, is un- 
doubtedly the same as T. rustica. 

It is smaller (6 to 7 millim. long), more slender, the aperture not 
so large, and the ribs less numerous (20 to 25), and the basal keel is 
more conspicuous than in T. valida, and is continuous with tlie large 
rib just behind the peristome, which gives the latter a duplicated 
appearance. The colour corneous or ruddy corneous. 

3. Truncatella ceylanica, Pfeiflfer. 

Truncatella ceylanica, Pfeiffer, Proc. Zool. Soc. 1S56 ; Mon. 
Auric. (Appendix) i. p. 186 ; H. Nevill, Enum. Pneum. Ceyl. 1871, 
p. 6 ; Tennent's Ceylon, i. p. 239. 

Truncatella teres", Vk\^er, Proc. Zool. Soc. 1856, p. 336; Mon. 
Auric. (Appendix) i. p. 188 ; Cox, Mon. Austr. Land-Shells, p. 92, 
pi. 15. figs. 9, 9a, 96 ; Paetel, Cat. Conch, p. 118. 

Truncatella semicostata, Montrouzier, Journ. de Conch. 1862, 
p. 243, pi. 9. fig. 10 ; Pfeiifer, Mon. Pneum. iii. p. 6 ; Gassies, 
Faune Nouv. Caled. p. 73, pi. 8. fig. 2 ; Paetel, Cat. Conch, p. 118 ; 
Schmeltz, Cat. Mus. Godeif. v. p. 104. 

T)-uncatella cerea, Gassies. 

Truncatella nitida, Gassies. 

We obtained many thousand specimens, in all stages of growth, 
near high-water mark at Ovalau Island. 

This species may be distinguished by its rather thin shining 
texture and more or less evanescent riblets, which are either well 
developed on all the whorls, or entirely absent except at the sutures, 
where they are reduced to plicate crenulations. Sometimes the crenu- 
lations disappear, so that the shell is perfectly smooth and polished. 
Some examples have the upper whorls ribbed, and the lower one 
smooth or crenulated at the suture and base. 

Having lately received from my esteemed correspondent, Mr. E. 
L. Layard, of New Caledonia, a lot of Truncatella ceylanica from 
Ceylon, together with T. teres from the Comoro Islands, and many 
examples of T. semicostata from New Caledonia, and after a critical 
comparison of the three species, I have failed to discover a single 
specific character to separate one from the other. Dr. Cox records 
T. teres from N.E. Australia. 

Through the courtesy of the Rev. Montrouzier of New Caledonia, 



IS87.] MOLLUSKS OF THE VITI ISLANDS. 301 

I have been enabled to examine typical specimens of T. cerea and 
T. nitidn labelled in Gassies's own handwriting, and do not hesitate 
to refer the former to the smooth crenulated and the latter to the 
smooth non-crenulated varieties of T. ceylanica. 

4. Truncatella granum, Garrett. 

Trunraiella granum, Gairett, Amer. Journ. Conch. 1872, p. 225 ; 
Schmeltz, Cat. Mus. GodefF. v. p. 105 ; PfeifFer, Mon. Pneum. iv, 
p. 16. 

This small species occurred to my notice in one locality only, the 
N.E. end of Taviuni, where it was found abundant beneath loose 
stones a little below high-water mark. 

Its small size (4| to 5 millim. long), uniform cinereous colour, 
4h persistent whorls, 20 to 25 riblets on the body-whorl, small ovate 
aperture, and conspicuous duplicated peristome will distinguish it 
from the preceding species. 

5. Truncatella avenacea, sp. nov. 

Shell small, rimate, cylindrical, corneous ; ribs rather small, erect, 
rounded, about the same width as their interspaces, 35 to 40 on the 
last whorl ; suture impressed ; whorls persistent, 4|, convex; base 
distinctly carinated ; aperture small, vertical, oval, a little less than 
a fourth the length of the shell ; peristome continuous, obtuse, 
slightly expanded, and duplicated by the continuation of the basal 
keel. Length 6, diam. 2| millim. 

A few examples found associated with T^ vitiana at Yanua Levu. 

It more nearly resembles the strongly ribbed T. semicostata than 
any other species inhabiting the group ; but may be distinguished 
from that species by its more numerous ribs, more convex whorls, 
smaller aperture, and more conspicuous basal keel and duplicated 
peristome. 

Genus Taheitia, H. & A. Adams. 

1. Taheitia funiculus (Mousson), 

Truncatella funiculus, Mousson, Journ. de Conch. 1870, p. 171 ; 
Paetel, Cat. Conch, p. 117; Pleiffer, Mon. Pneum. iv. p. 21. 

This species was discovered by Dr. Griiffe in the interior of Viti 
Levu. 

It is a slender, cylindrical, whitish horn-coloured species, with 6 
persistent convex whorls and a deep suture ; the ribs are sharp, ratiier 
remote, 12 to 16 in the body-whorl, all converging at the base. The 
aperture is small, vertical, semicircular. Peristome obtuse, expanded, 
continuous, and slightly porrected. Length 8, diam. 2 millim. 

2. Taheitia turricula (Mousson). 

Truncatella turricula, Mousson, Journ. de Conch. 1870, p. 196; 
Paetel, Cat. Conch, p. 117 ; Schmeltz, Cat. Mus. Godeff. v. p. 104 ; 
Pfeiffer, Mon. Pneum. iv. p. 20. 

Inhabits Mango Island, where it was discovered by Dr. Griiffe. 
I do not know this species. 

Proc. Zool. Soc— 1887, No. XXI. 21 



302 MR. A, GARRETT ON THE TERRESTRIAL [Mar. I , 

It is a long, slender, greyish-white species, with 5 persistent 
subconvex whorls and a deep suture ; the riblets, of which there are 1 8 
to 20 on the last whorl, are thin, sharp, and separated by rather wide 
interspaces. The vertical aperture is obtusely biangulately ovate, 
and the porrected peristome is continuous, expanded, and slightly 
reflected. Length 9 millim. 

3. Taheitia arcasiana (Crosse). 

Truncatella arcasiana, Crosse, Journ. de Conch. 18G8, p. 177, 
1870, p. 107, pi. 7. fig. 13; Mousson, Journ. de Conch. 1870, 
p. 196; Schmeitz, Cat. Mus. Godeff. v. p. 104; Pfeiffer, Mon. 
Pneum. iv. p. 20. 

Truncatella alternans, Mousson, in Cat. Mus. Godeff. iv. p. 7G- 

This species (which is unknown to me) was found by Dr. Griiffe in 
the interior of Vjti Levu. 

Mr. Crosse says it is an elongated, cylindrically-turreted, chalky, 
dull whitish species, 9 millim. long, with remote riblets, which are 
sometimes evanescent on the middle of the whorls. The persistent 
6 whorls are rather convex, the suture is impressed, and the aperture 
suboval. The peristome is continuous, porrected, and slightly 
expanded. 

4. Taheitia scalariformis (Reeve). 

Truncatella scalariformis. Reeve, Proc. Zool. Soc. 1842, p. 197; 
Conch. Syst. ii. pi. 182. fig. 6 ; Pfeiffer, Zeit. Malak. 1846, p. 186 ; 
Mon. Auric, i, (Appendix) p. 191 ; Jay, Cat. Shells, 1850, 
p. 252 ; KiJster, Mon. p. 15 ; H. & A. Adams, Gen. Moll. ii. p. 31 ; 
Schmeitz, Cat. Mus. Godeff. v. p. 104. 

Truncatella truncatula, var., Anton, Verz. Conch, p. 62. 
Truncatella arctecostata, Mousson, Journ.de Conch. 1869, p. 68, 
pi. 5. fig. 4, 1870, p. 195; Paetel, Ciit. Conch, p. 117; Schmeitz, 
■ Cat. Mus. Godeff. iv. p. 76 ; Pfeiffer, Mon. Pneum. iv. p. 19. 

Taheitia scalariformis, Pease, Proc. Zool. Soc. 1871, pp. 468, 
477; Journ. de Conch. 1871, p. 92; Martens & Langk. Don. 
Bismark. p. 60, pi. 4. fig. 1. 

Prof. Mousson's original description of T. arctecostata was drawn 
up from specimens of T. scalariformis collected by me at Anaa, 
Paumotu Islands. The same, or a very closely allied, species was 
subsequently discovered by Dr. Griiffe on Viti Levu, and referred by 
Mousson to T. arctecostata. I have not seen any Viti specimens. 

It is a thin, subpellucid, cylindrically-turreted, yellowish-white 
or corneous species, with 4 or 5 persistent convex whorls. The 
riblets are numerous (38 to 40 in the last whorl), very slightly 
arched, and converging at the base. The aperture is vertical, small, 
broadly ovate, and the peristome is thin, continuous, and expanded. 
Operculum typical. Length 6 millim. 

Genus Diplommatina, Benson. 
1. Diplommatina martensi, H. Adams. 
Diplommatina (Diancta) martensi, H. Adams, Proc. Zool. Soc. 



1887.] MOLLUSKS OF THE VITI ISLANDS. 303 

1866, p. 446, pi. 38, fig. 11 ; Mousson, Journ. de Conch. 1870, 
p. 186; Brazier, Journ. de Conch. 1870, p. 84; Schmeltz, Cat. 
Mus. Godeff. v. p. 102 ; Pfeiffer, Mou. Pueum. iv. p. 85. 

Diplommatina paradoxa, Crosse, Journ. de Co.icli. 18.57, p 449. 

Diplommatina austral'KB, Schmeltz (not of Benson), Cat. Mus. 
Godeff. iii. p. 30; Paetel, Cat. Conch, p. 118. 

Diplommatina macrostoma, " MSS.," Schmeltz, Cat. Mus. Godeff. 
iv, p. 7.T ; Paetel, Cat. Conch, p. 118. 

Diplommatina distorta, " MSS.," Schmeltz, I. c. p. 75 ; Paetel, 
/. c. p. 118. 

This species inhabits Viti Levu and Ovalau, 

It may be distinguished by its sinistral, distorted, ovate-couical 
form, cinereous or luteous horn-colour, 5| swollen whorls, the last one 
smaller than the preceding and ascending. The sculpture consists 
of fine, crowded, oblique, lamelliform striee, wliich become larger 
and more remote on the body-whorl. Aperture large, subvertical, 
and nearly circular. The peristome is continuous, expanded, and 
in adults the parietal wall is subplicate, JiCngth 3 millim. 

2. Diplommatina pomati^formis, Mousson. 
Diplommatina (Diancta) pomaticBformis, Mousson, Journ. de 

Couch. 1870, p. 180, pi, 8. fig. 2 ; Pfeiffer, Mon. Pneum. iv. p. 81, 
Discovered by Dr. Griiffe in the central portion of Viti Levu. 
A sinistral, inflated, costate-striated, cinereous species, with 6k 
rounded whorls, the last one smaller than the penultimate, scarcely 
ascending, and the striae coarser and more distant than on the pre- 
ceding whorls. The vertical aperture is circular, pale luteous, and 
the peristome is slightly reflected and duplicated. Length 5 millim. 

3. Diplommatina subregularis, Mousson. 

Diplommatina (Diancta) subregularis, Mousson, Journ. de Conch. 
1870, p. 181, pi. 8. fig. 3 ; Pfeiffer, Mon. Pneum. iv. p. 81, 

This species inhabits " Nagara," on the south coast of Viti Levu, 
where it was found by Dr. Griiffe. 

A sinistral, yellowish horn-coloured species, shaped like the pre- 
ceding, but not so much inflated and with the spire more regular. 
The sculpture consists of distant acute striae. Whorls 7, convex, 
the penultimate somewhat inflated, the last one small and ascending. 
The vertical aperture is quadrately circular, and the slightly expanded 
peristome is subduplicated. Length 3 millim, 

4. Diplommatina ascendens, Mousson. 

Diplommatina {^Diancta) ascendens, Mousson, Journ, de Conch. 
1870, p. 184, pi. 8. fig. 5 ; Pfeiffer, Mon. Pneum. iv. p, 82. 
Dr. Griiffe discovered this species on the island of Viti Levu, 
It is a sinistral species about 3g millim. long, of a flesh-white 
colour, acutely ovate form, with distant lamelliform riblets and 
rather deep suture. Whorls 5|, convex, rapidly increasing, the 
last one smaller than the penultimate and ascending. The rather 
large aperture is somewhat quadrate in shape and the peristome is 

21* 



304 MR. A. GAnRETT ON THE TERRESTRIAL [Mar. 1, 

expanded, duplicated, slightly sinuated, and the columella is obtusely 
nodulous. 

5. DiPLOMMATINA GODEFFROYANA, MoUSSOU. 

DipJommatina {Diancta) godeffroyana, Mousson, Journ. de Conch. 
1870, p. 182, pi. 8. tig. 4; Paetel, Cat. Conch, p. 118; Schmeltz, 
Cat. Mus. Godeff. v. p. 102 ; Pfeiffer, Mon. Pneum. iv. p. 82. 

Also discovered by Dr. GriifFe on the southern portion of Viti 
Leva. 

It is described as a sinistral, ovate, inflated, whitish horn-coloured 
species, with distant lamina-like strife, 6 whorls, the antepenultimate 
larger than the penultimate, and the latter larger than the body- 
whorl, which slightly ascends the preceding one. The subvertical 
aperture is circular ; the peristome is acute, shortly expanded, and 
slightly duplicated. Length 3g millim. 

6. DiPLOMMATINA TUBEROSA, MouSSOU. 

Diplommatina (Diancta) tuberusa, Mousson, Journ. de Conch. 
1870, p. 185; Paetel, Cat. Conch, p. 118; Schmeltz, Cat. Mus. 
Godeff. V. p. 102 ; Pfeiffer, Mon. Pneum. iv. p. 83. 

This is another of Dr. GrafFe's newly discovered species, which he 
found at Vaini Loba, on the south part of Viti Levu. 

It is described as a sinistral, rather thin, elongate-ovate, pale 
horn-coloured species, with distant lamelliform riblets, 6 rounded 
whorls, the penultimate retracted and compressed in front, and the 
sides inflated. The last whorl is small, attenuated, and slightly 
ascending. The aperture is subcircular, and the peristome is slightly 
reflexed. Length 3-4 millim. 

7. DiPLOMMATINA QUADRATA, MoUSSOU. 

Diplommatina (Diancia) guadraia, Mousson, Journ. de Conch. 
1870, p. 187, pi. 8. fig. 1 ; Pfeiffer, Mon. Pneum. iv. p. 83. 

Also found by Dr. Graffe at Viti Levu. 

A sinistral, ovate, pale-yellowish species, with crowded costulate 
striae and convexly conical spire. Whorls 5, rapidly increasing, 
convex, the fourth one subinflated, the penultimate swollen on the 
back and retracted on the front, the last one attenuated, compressed 
at the base, and rapidly ascending. The subquadrate aperture is 
subpatulous, and the peristome is expanded. Length 4| millim. 

8. DiPLOMMATINA TAViENsis, Liardct. 

Diplommatina taviensis, Liardet, Proc. Zool. Soc. 1876, p. 101, 
pi. .O. figs. 9, 9«. 

" Shell with the penultimate whorl contracted in front, leaving the 
previous one and hp of the aperture joining regularly costated ; 
lip double ; aperture circular and entire." 

" Animal with two tentacles, short and cylindrical, with an active 
arched motion, as in Helicina. Eyes situated at the base of the 
tentacles inside." 

" Hab. Taviuni, Fiji." {Liardet.) 



1887.] MOLLUSKS OF THE VITI ISLANDS. 305 

Genus MoussoNiA, Semper. 

1. MoussoNiA FuscuLA, Mousson. 

Diphmmatina (JSIoussonia) fuscula, Mousson, Journ. de Conch. 
1H70, p. 188, pi. 8. fig. 9 ; Pfeiffer, Men. Pneum. iv. p. 93. 

Moussonia fuscula, Paetel, Cat. Concb. p. 102. 

Tlie type was found by Dr. Griiffe on Oneata, one of the Windward 
Islands. Prof. Mousson mentions a var. viliana from Viti Levu. 

It is a minute, dextral, abbreviately turreted species of a brownish 
horn-colour, with fine striae and 7 rounded whorls, of which the 
penultimate is the larger. The body-whorl is rather slender, 
rounded, and ascending. The aperture is circular, and the columella 
bears an obtuse plait or tooth. Peristome expanded. Length 
2 millim. 

Genus Ostodes, Gould. 

1. Ostodes diatretus (Gould). 

Cyclostoma diatretum, Gould, Proc. Bost, Soc. Nat. Hist. 1847, 
p. 205 ; Expl. Exp., Shells, p. 105, fig. 124. 

Cyclotus diatretus, Pfeifter, Consp. Cyclos. no 22 ; Mon. Pneum. 
i. p. 33 ; Gray, Cat. Phan. p. 18. 

Cyclophorus (Ostodes) diatretus, var. intercostata, Mousson, 
Journ. de Conch. 1870, p. 179. 

Dr. Gould's type specimens of this rare species were found on the 
west end of Vanua Levu. I obtained two dead specimens at Vanua 
Balavo, one of the Windward Islands, where Dr. GriifPe also found 
a single dead example, and one on Oneata, which Prof. Mousson 
characterized as var. irdercostata. 

This species is readily distinguished by its depressed form, wide 
open umbilicus, rude spiral, unequal, elevated lines, whitish horn- 
colour, rounded aperture, and simple peristome. Diam. 12 millim. 

It is very closely related to the New-Caledoniau O. bocayeanus, 
but is smaller and more depressed. 

Whilst searching in the mountains on the west end of Vanua 
Levu I found a very much weathered Ostodes twice the size of 
Gould's s[)ecies, and subsequently found a similar specimen on the 
beach at Kioa Island. The specimens are in the Museum Godeffroy 
in Hamburg, It is undoubtedly an undescribed species. 

2. Ostodes liberatus, Mousson. 

Ostodes liberatus, Mousson, MS., Mus. Godeffroy, 1885. 

Shell widely umbilicated, depressed, whitish horn-colour ; spire 
slightly elevated; apex prominent; suture deeply impressed; 
whorls 4, convex, transversely rudely striated, undulated, and 
sculptured with numerous elevated spiral lines, larger, more promi- 
nent, and crenulated on and above the rounded periphery; umbilicus 
very wide, showing all the volutions to the apex ; aperture circular, 
nearly vertical ; peristome thin, continuous, straight. 

Major diam. 10, height 4 millim. 

Viti Levu. 



306 MR. A. GARRETT ON THE TERRESTRIAL [Mar. I, 

I received four examples of this singular species from the Museum 
Godeffroy. One specimen has the last whorl separated from the 
penultimate a distance of 4 millim. It may he distinguished by 
its depressed form, undulated whorls, and crenulated spiral lines. 

3. OsTODES STRiCTUs, Mousson. 

Ostorles strictus, Mousson, MS., Museum Godeffroy, 188.5. 

Shell umbilicated, depressed, turbinate, solid, rugose, decorticated, 
cinereous, sometimes with a ruddy tinge on the last whorl ; spire 
depressedly conoid, apex exserted ; suture impressed ; whorls 5, con- 
vex, transversely rugosely wrinkled, closely lineated with spiral 
elevated lines, becoming evanescent on the rounded body-whorl ; 
umbilicus wide, freely exhibiting all the whorls, spirally lineated 
with raised lines, and the margins slightly angulated ; aperture 
oblique, subcircular ; peristome straight, simple, nearly continuous, 
briefly joined to the body-whorl. 

Major diam. 13, height 7 millim. 

Vatu Lale. 

Three examples received from the Godeffroy Museum. It is very 
closely related to Gould's 0. strigatus, a Samoa species, and, excepting 
in size, can scarcely be distinguished from the New-Caledonian 
O. bocageanus. 

Genus Pupina, Vignard. 
1. Pupina viTiENSis, Garrett. 

Pupina vitiensis, Garrett, Proc. Phil. Acad. Nat. Sci. 1873, p. 233, 
pi. 3. fig. 62; Schmeltz, Cat. Mus. Godeff. vi. pp. 83, 104. 

A somewhat rare species, found beneath damp decaying leaves at 
Gomea Island. Mr. Liardet records a species of Pupina as occur- 
ring on Taviuni, which is probably the same as the Gomea shell. 
Schmeltz erroneously assigns it to Kandavu. 

A brilliant, highly polished, oblong, whitish corneous species, with 
slightly swollen spire, the left side more convex than the right, and 
the columella with a tongue-like projection forming a deep notch. 
An ohtuse plait on the upper part of the parietal wall. Length 7 
millim. 

Genus Omphalotropis, Pfeiffer. 

1. Omphalotropis moussoni. Pease. 

Omphalotropis ovata, Mousson (not of Pease), Journ. de Conch. 
1865, p. 198, pi. 14. fig. 10; Paetel, Cat. Conch, p. 124 ; Schmeltz, 
Cat. Mus. Godeff. iv. p. 75. 

Omphalotropis moussoni. Pease, Journ. de Conch. 1869, p. 147; 
Schmeltz, Cat. Mus. Godeff. v. p. 101. 

Realia {Omphalotropis) moussoni, Mousson, Journ. de Conch. 
1870, p. 194, 1871, p. 27 ; Pfeiffer, Mon. Pneum. iv. p. 224. 

I found a few examples of this species at Vaima Balavo, where 
Dr. GriifPe discovered the type specimens. The Doctor subse- 
quently found it on Yiti Levu, Ticombia, and at Tongatabu, one of 



1887.] MOLLUSKS OF THE VITI ISLANDS. 307 

the Tonga group. All the sj)ecies of this genus live beneath decay- 
ing vegetation. 

A smooth, ovate-veutricose, yellowish horn-coloured species, 
with two faint transverse zones, and six subiuflated whorls, the last 
one perforated and slightly keeled close to the perforation. Length 
3| millim. 

2. Omphalotropis parva, Mousson. 

Omphalofropis parva, jNIousson, Journ. de Conch. 186.5, p. 199 ; 
Pease, Journ. de Conch. 1869, p. 147 ; Paetel, Cat. Conch, p. 124; 
Schmeltz, Cat. Mus. Godeff. v. p. 101. 

Realia (Omphalotropis) parva, Mousson, Journ. de Conch. 1871, 
p. 28 ; Pfeiffer, Mon. Pneuni. iv. p. 224. 

Realia Icevis, Baird, in Brenchley's Cruise of Curafoa (ex Schmeltz, 
in C. M.G. V. p. 101). 

Omphalotropis vitiensis, Liardet, Proc. Zool. Soc. 1876, p. 101, 
pi. 5. figs. 11, 11a. 

This small species is not only generally diffused throughout the 
group, but occurs also in the Tonga and Ellis group of islands. 
Dr. i3aird gives "Samoa" as habitat; neither Griiffe nor myself 
detected it in that group. 

A small, smooth, ovate-conical species, 4| millim. long, of a pale 
corneous, yellowish, or violaceous horn-colour, with 6 convex whorls, 
and a strong basal keel contiguous to the perforation. 

3. Omphalotropis ingens, Mousson. 

Realia {Omphalotropis) ingens, Mousson, Journ. de Conch. 1870, 
p. 189 ; Pfeiffer, Mon. Pueum. iv. p. 227. 

This species was found by Dr. Graffe in Oneata, where it occurred 
in a semi-fossil condition. 

An acutely-ovate species, with longitudinal riblets and 7 flattened 
whorls, the last with a basal filiform keel. Length 7 millim. 

4. Omphalotropis LONGULA, Mousson. 

Realia {Omphalotropis) longula, Mousson, Journ. de Conch. 
1870, p. 193 ; Pfeiffer, Mon. Pneum. iv. p. 233. 

Omphalotropis longula, Paetel, Cat. Conch, p. 124. 

Inhabits Ticombia, one of the Windward Islands. 

This species may be characterized by its smooth, rather thin, sub- 
pellucid, conically-turreted form, 7 sliglitly convex whorls, the last 
one rounded and filocarinate at the base. The aperture is oval, and 
the peristome slightly expanded. Length 6 millim. 

5. Omphalotropis circumlineata, Mousson. 

Realia {Omphalotropis) circumlineata, Mousson, Journ. de Conch. 
1870, p. 191, pi. 7. fig. 11 ; Pfeiffer, Mon. Pneum. iv. p. 230. 

Omphalotropis circumlineata, Paetel, Cat. Conch, p. 124. 

Garrettia 1 circumlineata, Schmeltz, Cat. Mus. Godeff. v. p. 100. 

This interesting species was discovered by Dr. Graffe on Vanua 
Balavo and Viti Levu. 



308 MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

A tliiii, turbinate, conical, reddish horn-coloured species, with 6 
angulate whorls which are spirally lineated with elevated lines, two 
on the body and one on the whorls of the spire so much larger than 
the others as to give the former a biangular, and the latter an angu- 
lar outline. The basal keel is small. Length 5| millim. 

]\Iousson's figure is not very characteristic. The whorls are too 
much rounded, and do not exhibit the large spiral lines which modify 
the outlines of the shell. 

Mr. Schmeltz refers it with a doubt to the genus Garrettia 
{ = Diadema, Pease). An examination of the operculum would de- 
cide the question of its generic rank. 

6. Omphalotropis costulata, Mousson. 

Eealia {Omphalotropis) costulata, Mousson, Journ. de Conch. 
1870, p. 190, pi. 7. fig. 10 ; Pfeiffer, Mon. Pneum. iv. p. 233. 

Omphalotropis costulata, Paetel, Cat. Conch, p. 124. 

A few examples found beneath dead leaves at Vanua Balavo, 
where Dr. Grjiffe obtained the type specimens. 

A small, ovate, whitish horn-coloured species, 5| millim. long, 
with 6 convex whorls, and furnished with small, longitudinal, 
crowded riblets. The perforated base has a well-defined keel, and 
there is sometimes faint indication of a peripheral keel. 

7. Omphalotropis subsoltjta, Mousson. 

Realia {Omphalotropis) subsohda, Mousson, Journ. de Conch. 
1870, p. 192, pi. 7. fig. 12; Pfeiffer, Mon. Pneum. iv. p. 219. 

Omphalotropis subsulnta, Paetel, Cat. Conch, p. 124 ; Schmeltz, 
Cat. Mus. Godeff. v. p. 100. 

Discovered by Dr. Griiffe at Oneata, one of the Windward Islands. 

A smooth, pale horn-coloured, turreted species, with 7| convex 
whorls; the List one, which is slightly separated from the penul- 
timate, is usually furnished with a filiform carina a little below the 
periphery. A similar remote keel circumscribes the basal perfora- 
tion. The vertical aperture is about one fifth the length of the 
shell. Peristome porrected, continuous, and slightly patulous at the 
base. Length 10 millim. 

8. Omphalotropis zebriolata, Mousson. 

Omphalotropis zebriolata, Mousson, Journ. de Conch. 1865, 
p. 181, pi. 14. fig. 11, 1870, p. 193, 1873, p. 108; Pease, Journ. 
de Conch. 1869, p. 145; Proc. Zool. Soc. 1871, p. 476; Paetel, 
Cat. Conch, p. 124; Schmeltz, Cat. Mus. Godeff. v. p. 101. 

Realia {Omphalotropis) zebriolata, Mousson, Journ. de Conch. 
1870, p. 193, 1871, p. 27; Pfeiffer, Mon. Pneum. iv. p. 225. 

Omphalotropis perforata, Mousson, Journ. de Conch. 1865, 
p. 182, pi. 14. fig. 12; Pease, Journ. de Conch. 1869, p. 145; 
Proc. Zool. Soc. 1871, p. 476; Paetel, Car. Couch, p. 124; 
Schmeltz, Cat. Mus. Godeff. v. p. 101. 



1887.] MOLLUSKS OF THE VITI ISLANDS. 309 

Realia (Omphalotropis) ■perforata, Mousson, Journ. de Conch. 
1871, p. 27; Pfeitfer, Mon. Pneum. iv. p. 222. 

The type specimens of O. zebriolata and O. perforata were found by 
Dr. Griiffe at " Uvea" or Wallis Island, one of the northern islands of 
the Tonga group. Having personally collected hundreds of examples 
of both forms in the same locality, and as they gradually intergrade, 
I have united the two species. Dr. Gr;itfe also detected it on the 
neighbouring island of Futuna and on the low coral-islands of Ellis 
group. He likewise obtained the form O. zebriolata at Kanathia, Viti 
Islands. Mr. Pease wrongly assigns it to Samoa. 

It may be distinguished by its ovate-conical form, smooth surface, 
fissured base, and variable colour — rose-red, pale luteous, violaceous 
brown, often pale banded at the periphery; sometimes with pale dots 
or longitudinal flexuous or zigzag lines. The basal keel is contiguous 
to the umbilical fissure. The pyiifbrm aperture is usually concolor 
and the glazed parietal wall is frequently reddish brown. Length 
5 to 7 millim. 

It is shaped like the well-known O. huahinensis. 

9. Omphalotropis rosea (Gould). 

Cyclostonia roseum, Gould, Proc. Bost. Soc. Nat. Hist. 1847, 
p. 2U5 ; Expl. Exp., Siiells, p. lO.i, fig. 121 ; Petit, Journ. de Conch. 
18.50, p. 47. 

Omphalotropis rosea, Pffiffer, Proc. Zool. Soc. 1854, p. 307 ; 
Consp, Cyclos. no. 47; Mon. Pneum. i. p. 308, iii. p. 176; H. & 
A. Adams, Gen. ^loll. ii. \\. 300. 

Hydrocena rosea, Pfeiffer, Mon. Pneum. ii. p. 162. 

Assiminea rosea. Martens, Ann. & Mag. Nat. Hist. 1866, xvii. 
p. 206. 

Through the kindness of Mr. E. L. Layard, I have received an 
adolescent example of Gould's 0. rosea ; but, unfortunately, the 
kind donor cannot give me any information in regard to the exact 
locality except the comprehensive one " Fiji." I am inclined to 
believe it inhabits the sm.all Windward Islands. 

It may be distinguished by its large size, elongate-conical shape, 
rather solid texture, 6—7 whorls, and rose or luteous horn-colour. 
The conspicuous basal keel is close to the slightly perforated 
umbilicus. Length S-U, diam. 41-5 millim. 

10. Omphalotropis bifilaris, Mousson. 

Omplialf.tropis hifiluris, Mousson, Journ. de Conch. 1865, p. 183 ; 
Pease, Journ. de Conch. 1869, p. 146; Proc. Zool. Soc. 1871, 
p. 476 ; Paetel, Cat. Conch, p. 124 ; Schmeltz, Gat. Mus. Godeff. v. 
p. 101. 

Benlia (Omphalotropis) bifilaris,'Moa&son, Journ. de Conch. 1869, 
p. 353, 1870, p. 194 (var. anc/usta), 1871, p. 29; Pfeiffer, Mon. 
Pneum. iv. p. 232. 

Obtained by Dr. Griiffe at Kanathia, and the var. angusta in the 
interior of VitiLevu. The latter has a smaller basal fissu.e, and the 



310 MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

two caiiiiations are evanescent. It is probably a distinct species. 
The tj'pe is also distributed throughout the group, and occurs at 
"WaUis Island. 

This species may be recognized by its conical form, pale horn- 
colour beneath a light brownish epidermis, 6 convex whorls, incised 
suture, and filocarinate periphery. The basal perforation is margined 
by a rather distant keel. Length 6 millim. 

] 1 . OiMPHALOTROPIS LAYARDIANA, Sp. UOV. 

Realea (Ompkalotropis) rosea, Mousson (not of Gould), Journ. de 
Conch. 1870, p. 192. 

Omphalotropis bythinaformis, " MSS.," Paetel, Cat. Conch, 
p. 124 (ex Schmeltz in Cat. Mus. Godeff. iv. p. 74). 

I obtained several specimens of this species at Vanua Balavo, one 
of ihe Windward Islands. It may be described as follows : — 

Shell umbilicated, rather solid, ovate-conic, smooth, scarcely 
shining, uniform corneous ; spire rather short, convexly-conical, apex 
obtuse ; suture slightly incised ; whorls 6-7, convex, narrowly 
tabulated, last one rounded ; umbilicus large ; basal keel large, and 
distant from the umbilical opening; aperture abbreviately ovate, 
angular po.-teriorly and rounded in front ; peristome obtuse, margins 
nearly or quite continuous. 

Length 6, diam. 4 millim. 

It differs from O. rosea in its smaller size, more abbreviate form, 
more swollen whorls, large umbilicus, and remote keel. 

Genus Lagochexlus, Blanford. 
1. Lagocheilus hispidus, Liardet. 

Lagocheiius hispidus., Liardet, Proc. Zool. Soc. 1876, p. 101, 
pi. 5. figs. 10, 10a. 

"Shell small, bulimoid, hispid, of a brown colour; whorls 5^, 
spirally costate: aperture circular. 

" Hab. Gomia, Fiji." {Liardet.) 

The existence of this East-Indian genus in the Viti group is 
remarkable. So far as known, it has not been detected in any other 
part of the Pacific. 

Genus Helicina, Lamarck. 
1. Helicina TECTiFORMis, Mousson. 

Helicina tectiformis, Mousson, Journ. de Conch. 1870, p. 199, 
pi. 8. fig. 7 ; Paetel, Cat. Conch, p. 126 ; {Trochatella) Brazier, 
Proc. Zool. Soc. 1871, p. 322 ; Schmeltz, Cat. Mus. Godeff. v. 
p. 98 ; Pfeiffer, Mon. Pneum. iv. p. 250. 

Helicina mangoensis, Sowerby, Proc. Zool. Soc. 1870, p. 250. 

This fine species appears to be peculiar to the small island of 
Mango, where it is very abundant on coralline rocks. 

x\ depressedly-conical, acutely carinated species of a uniform white 
colour, with a sulphur-yellow spire ; frequently greyish, more or less 



lyH/.] MOLLUSKS OF THE VITl ISLANDS. 311 

tinged with red. The sculpture consists of rather coarse, elevated, 
oblique strife of growth, aud closely-set spiral raised lines. The very 
oblique aperture is subtriangular, and the obtuse peristome is more 
or less expanded. There are 5 flat whorls, the last one with a 
prominent compressed acute keel. Diam. 12 millim. 

2. Helicina sempert, Mousson. 

Helicina semperi, Mousson, Journ. de Conch. 1870, p. 201, pi. 8. 
fig. S; Paetel, Cat. Conch, p. 126 ; Schmeltz, Cat. Mus. Godeff. v. 
p. 98 (as of Graflfe) ; Pfeiffer, Mon. Pneum. iv. p. 2/8. 

Obtained by Dr. Graflfe at Oneata. 

A solid, smooth, somewhat lenticular, shining species with five 
flattened whorls, the last one rounded at the periphery, and tlie 
colour variable — white, yellow, reddish, frequently with reddish 
zigzag strigations. Peristome obtusely thickened, and the basal 
callus white. Diam. 10 millim.. 

3. Helicina interna, Mousson. 

Helicina interna, Mousson, Journ. de Conch. 1870, p. 201, pi. 8. 
fig. 6; 1871, p. 24; Paetel. Cat. Conch, p. 125; Schmeltz, Cat. 
Mus. Godefi". v. p. 99 ; Pfeiffer, Mon. Pneum. iv. p. 248. 

Found by Dr. Grafte in the interior of Viti Levu and at Mango 
Island. It is also recorded from the Tonga group. 

A turbinately-conical species of a luiiform white or yellowish 
colour, with or without a spiral reddish-brown zone, and regular 
conical spire. Whorls .5, slightly convex, the last one rounded or 
obtusely angulated. Peristome acute. Diam. 9 millim. 

4. Helicina gomeaensis, Garrett. 

Helicina qomeacnsis, Garrett, Proc. Phil. Acad. Nat. Sci. 1673, 
p. 233, pi. 3. fig. 13. 

On the foliage of bushes at Gomea Island. 

A depressedly trochiform, somewhat shining species, with spiral 
impressed striae. Colour light straw-yellow, rarely with two brownish- 
red zones. Whorls 5, somewhat convex, last one slightly angular 
on the periphery. The white peristome is broadly expanded. Diam. 
10 millim. 

5. Helicina pallida, Goiild. 

Helicina pallida, Gould, Proc. Best. Soc. Nat. Hist. 1847, 
p. 202 ; Expl. Exp., Shells, p. 96, fig. 113 ; Pfeiffer, Mon. Pneum. i. 
p. 396 ; Gray. Cat. Phan. p. 290; {Pachystoma) H. & A. Adams, 
Gen. Moll. ii. p. 303; Mousson, Journ. de Conch. 1865, p. 197, 
1870, p. 200; Paetel, Cat. Conch, p. 125; Schmeltz, Cat. Mus. 
Godeff. v. p. 74. 

I found a few examples of this species at Vanua Levu, Kioa, and 
Vanua Balavo. On foliage. 

A cinereous or pale yellowish-white species, more depressed than 
the preceding one, the whorls flatter, the last one angular, the 



312 MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

spiral lines more indistitict, and the periphery with a compressed keel. 
The peristome is thinner and not so much expanded. Diam. 
9 millim. 

6. Helicina beryllina, Gould. 

Helicina heryllinu, Gould, Proc. Bost. Soc. Nat. Hist. 184/, 
p. 202 ; Expl. Exp., Shells, p. 95, fig. 1 1 1 ; Pfeiffer, Mon. Pneum. i. 
p. 354; Gray, Cat. Phan. p. 25(i ; {Mesa) H. & A. Adams, Gen. 
Moll. ii. p. 304 ; Mousson, Jcurn. de (^oiich. 1865, p. 197, 18()9, 
p. 357 (var. /aivWff), 1870, p. 200; Paetel, Cat. Conch, p. 125; 
Pease, Proc. Zool. Soc. 1871, p. 476 ; Schmeltz, Cat. Mas. Godeff. 
V. p. 98. 

I obtained numerous examples of this species at Vanua Balavo, 
where it occurs on the trunks of trees. Dr. Graffe found it at 
Kanathia and Oneata. Prof. ]Mousson has described ihexax.Jiavida 
from Samoa. 

About the same size but thinner than H. seniperi, and the spire 
is more conical, and the last whorl more depressed, so much so as to 
give the jjeriphery an obtusely angular appearance. The peristome 
is thin. Colour white or pale greenish yellow, frequently with a 
dorsal spiral red zone ; basal callus greenish yellow. 

7. Helicina fulgora, Gould. 

Helicina fulgora, Gould, Proc. Bost. Soc. Nat. Hist. 1847, 
p. 201 ; Ex])!. Ex])., Shells, p. 97, fig. 106 ; Pfeiffer, Mon. Pneum. 
i. p. 401 ; Gray, Cat. Phan. ]>. 293 ; H. & A. Adams, Gen. Moll. ii. 
p. 302 ; Mousson, Journ. de Conch. 18(15,]). 178, 1809, p. 356, 
1870, p. 198 (var. eocpansa), 1871, p. 25 (var. climinutu) ; Pease, 
Proc. Zuol. Soc. 1871, l'. 476 ; Paetel, Cat. Conch, p. 125 ; Schmeltz, 
Cat. Mus. Godeff. v. p. 98. 

Dr. Gould's type specimens were obtained at Manna and Upolu, 
Samoa Islands. Mouston's variety expansa was found by Dr. Giaffe 
at Kanathia and Viti Levu. He also described the small var. 
dimiuuta Ironi the Tonga Islands. Pfeiffer erroneously cites the 
Sandwich group as one of its localities. 

This species, which lives on the ground in forests, may be dis- 
tinguished by its thin, smooth, shining thell, convexly conoid spire, 
angular and conspicuously carinated body-whorl. The peristome 
is thin and expanded, forming an angle at the point of union with 
the columella. Colour pale corneous, white, or light sulphur-yellow, 
■with longitudinal flexuous reddish or white narrow stripes. Diam. 
4 to 9 millim. 

8. Helicina musiva, Gould. 

Helicina musiva, Gould, Proc. Bost. Soc. Nat. Hist. 1847, p. 201 ; 
Expl. Exp., Shells, p. 98, fig. 107; Pfeiffer, Mon. Pneum. i. 
p. 368 ; Gray, Cat. Phan. p. 25'9 ; H. & A. Adams, Gen. Moll. ii. 
p. 302; Mousson, Journ. de Conch. 1865, p. 178 (var. uveana), 
1869, p. 357, 1870, p. 202 (var. vitiana et sulcarinata), 1871, 



18*^7.] MOLtUSKS OF THE VITI ISLANDS. 313 

p. 25, 1873, p. 107 (var. rotundata); Pease, Proc. Zool. Soc. 18/1, 
p. 476 ; Paetel, Cat. Conch, p. 125 {musica in err.). 

This variable species is abundant beneath decaying vegetation in 
the lowlands near the sea-shore, and is generally diffused throughout 
the group. It is also common in similar stations in the Tonga and 
Samoa Islands. Prof. Mousson's var. rotundata was obtained at 
the low coral-islands of the Ellis group. 

The shape varies from depressed globose to sublenticular, and in 
size it varies from 3 to 5 millim. in diameter. The usual colour is 
white, corneous, or pale yellowish horn-colour, with radiating reddisli- 
chestnut, more or less zigzagged or undulating stripes ; it is rarely 
unicolor, and sometimes ligiht chestnut. The periphery is rounded 
or subangulated, and the peristome slightly expanded. 

H. oceanica. Pease, inhabiting the Kingsniill group, is probably 
a variety of H. musiva. 

9. Helicina articulata, Pfeiffer. 

HeHcina articulata, Pfeiffer, Proc. Zool. Soc. 1854, p. 53 ; Mon. 
Pneum. ii. p. 191 ; Malak. Blat. 1854, p. 103. 

A few examples found beneath decaying leaves near the sea-shore, 
on the west end of Vaniia Levu. 

Pfeiffer mentions "Tanna, New Hebrides," as the habitat of 
H. articulata. 

Very closely related to the depressed subangulated forms of 
H. musiva, and the colour and markings are nearly or quite similar 
in the two species. The two articulated white and chestnut bands 
at the suture and periphery are not constant. The filocarinated 
body-whorl is its most prominent character. Diam. 5-6 millim. 

10. Helicina pohliana, sp. nov. 

Helicina miniata, "Lesson" (not of Lesson in Voy. ('oquille). 
Museum Godeffroy, 1885. 

Shell solid, depressedly-conoid, smooth, shining, fulvous or luteous, 
with very faint oblique striae and distant irregular microscopicfil 
impressed spiral lines ; spire convexly conoid ; whorls 5, flatly 
convex, last one obtusely carinate ; keel white, compressed beneath ; 
aperture small, diagonal, semioval ; peristome very thick, labiate 
within ; basal callus white or pale yellowish. 

Viti Levu. 

I received this species from the Museum Grodeffroy, labelled 
" H. miniata. Lesson." Lesson's species is peculiar to the island of 
Bolabola, one of the Society group. 

Its solid texture, yellow colour, obtuse white keel, and thick 
obtuse peristome will readily distinguish it from any other Viti 
species. 

11. Helicina iNcisA, Mousson. 

Helicina incisa, Mousson, MS., Museum Godeffroy, 1 885. 

Shell minute, depressed-conoid, very faintly striated by lines of 



314 MR. A. GARRETT ON THE TERRESTRIAL [Mar. 1, 

growth, and spirally lineated with fine impressed lines; colour 
yellowish, corneous, with or without radiating reddish spots mid 
bands ; spire conoid ; suture linearly impressed ; whorls 4|-5, 
flatly convex, slowly and regularly increasing, the last one depressed, 
subangulated on the periphery ; base convex ; aperture oblique. 
Innately oval ; peristome expanded, slightly obtuse. 

Diam. 4 milhm. 

Ouo Island. 

Several examples received from the Museum Godeffroy. Possibly 
only a variety of H. musiva with fine spiral impressed lines. 

Genus Georissa, Blanford. 

1. Georissa juvenilis (Mousson). 

Diancta juvenilis, Mousson, MS., Mus. Godeffroy, 1885. 

" Viti Levu." 

Shell imperforate, ovate-conical, rather solid, corneous ; spire 
conical, apex obtuse ; suture deeply impressed ; whorls 5, strongly 
convex, longitudinally obliquely plicately ribbed ; ribs rude, irregular, 
becoming more crowded and smaller near the peristome ; aperture 
suborbicular ; peristome simple, straight; columellar region thickened 
with callus. Length 2| millim. Several examples were received 
from the Museum Godeffroy. The oblique longitudinal riblets will 
at once determine this from any other Polynesian species. 

2. Georissa parta (Pease). 

Cydostoma parvuvi, Pease, Proc. Zool. Soc. 1864, p. 674. 

Clwndrella j^arva. Pease, Proc. Zool. Soc. 1871, pp. 465, 476; 
Pfeiffer, Mon. Pneum. iv. p. 294 ; Garrett, Journ. Acad. Nat. Sci. 
Philad. 1884, p. 106, pi. 3. fig. 41. 

Ilydrocena insularis, " Crosse," Mousson, in Museum Godeffroy, 
1885. 

Examples of this species received from the Museum Godeffroy, 
labelled '' Hydrocena insularis, Crosse, Viti Levu," do not differ from 
Pease's Chondrella parva inhabiting the Society Isles. It is shaped 
like G. juvenilis, but is a little larger, and the whole surface is 
smooth and somewhat shining. 

Genus Assiminea, Leach. 

1. Assiminea nitida. Pease. 

Hydrocena nitida. Pease, Proc. Zool. Soc. 1864, p. 674. 

Assiminea 7iitida, Pease, Journ. de Conch. 1869, p. 165, pi. 7. 
fig. 11; Proc. Zool. Soc. 1871, p. 476; Schmeltz, Cat. Mus. 
Godeff. V. p. 103 ; Garrett, Proc. Phil. Acad. Nat. Sci. 1879, p. 29 ; 
Journ. Phil. Acad. Nat. Sci. 1881, p. 408, 1884, p. 107. 

? Realia nitida, Pfeiffer, Mon. Pneum. iii. p. 202. 

Hydrocena parvula, Mousson, Journ. de Conch. 1865, p. 184, 
1873, p. 108. 

Omphalotropis parvula. Pease, Journ. de Conch. 1869, p. 155 
Proc. Zool. Soc. 1871, p. 476 ; Paelel, Cat. Ccnch. p. 124. 



1887.] MOLLUSKS OF THE VITI ISLANDS, 315 

Assiminea parvula, Pease, Proc. Zool. Soc. 1871, p. 470 ; 
Schmeltz, Cat. Mus. Godeff. v. p. 103. 

Eealia parvula, Pfeiffer, Mon. Pneum. iii. p. 213. 

Assiminea liicida. Pease, Jourii. de Conch. 1869, p. ICG, pi. 7. 
fig. 10 ; Proc. Zool. Soc. 1871, p. 476. 

Assiminea ovata, " Pease," Schmeltz, Cat. Mus. Godeff. v. p. 103. 

Hydrocena pygmcea, Gassies, Journ. de Conch. 1867, p. 6.3. 

Assinmiea pi/gmcea, Pease, Journ. de Couch. 1869, p. 165. 

? Realia ptjgmcea, Pfeiffer, Mon. Pneum. iv. p. 214. 

Hydrocena sirnilis, Baird, iu Cruise of the ' Curacjoa.' 

This small species is generally distributed throughout all the 
groups from the Paumotus to the Viti Islands and New Caledonia, 
and ranges from near the sea-shore to about 2000 feet above the sea- 
level. They are found beneath decaying leaves, under stones and 
dead wood. 

It may be recognized by its small size, 2|-4 millim. long, smooth, 
shining surface, ovate-conical form, light or dark corneous colour, 
rarely with a faint transverse band on the last whorl. 

2. Assiminea brevissima (Mousson). 

Hydrocena brevissima, Mousson, Journ. de Conch. 1870, p. 194. 
Found by Dr. Graffe at Vanua Balavo. 

A minute, broadly conical, thin, pellucid, shining, pale reddish 
horn-coloured species vpith 3g whorls. Length 1| millim. 

3. Assiminea fischeriana (Gassies). 

Hydrocena fischeriana, Gassies, Faune Nouv. Caled. 1863, p. 115, 
pi. 7. fig. 18. 

Realia fischeriana, Pfeiffer, Mon. Pneum. iv. p. 421. 

Assiminea vitiensis, Garrett, Amer. Journ. Conch. 1872, p. 225, 
pi. 10. fig. 14. 

Abundant on the margins of mangrove-swamps. It also occurs 
at New Caledonia. 

Shaped like A. nitida, but larger, darker coloured, with or without 
one or two pale bands on the body-whorl. 



The following species, though quoted as Viti shells, have been 
found neitlier by Dr. Graffe nor by myself. Their existence in the 
group certainly needs confirmation. 

Parmella planata, H. Adams, Proc. Zool. Soc. 1867, p. o08, 
pi. 19. fig. 20. 

" Habitat Fiji Islands." (//. Adams.) 

Nanina SCORPIO, Gould, Expl. Exp., Shells, p. 33, fig. 67. 

" Ft ejee Islands." {Gould.) 

Not identified by any subsequent author. It has been referred 
to the genus Helicarion. 



31G MB. E. \. SMITH ON SHELLS [Mar. 1, 

PupiNA ADAMsiANA, Cfosse, Joum, dc Conch. 1871, p. 330 ; 
1872, p. 60, pi. 2. fig. 6. 

"VanuaLevu." (Crosse.) 

Mr. Crosse cites the locality on the authority of a London dealer. 
The species is closely allied to if not identical with Hargravesia 
polita, a Solomon-Island species. 

Helicina lens, Lea, Observ. i. p. 161, pi. 19. fig. 56. 

" Feejee Islands." {Lea.) 

Perhaps a unicoloured variety of JI. fulgora, Gld. 

Helix leucolena, Crosse, Joum. de Conch. 1867, p. 447 ; 
1868, p. 171, pi. 6. fig. 6. 

" Vanua Levu, Viti." (Crosse.) 

Mr. Crosse, who obtained the type specimen from a London 
dealer, was informed it came from Vanua Levu. The type is foreign 
to the group. 

Helix semirufa, Albers, Die Hel. p. 106. 
" Habitat in insulis Fidschi." (Albers.) 
Most certainly foreign to the group. 

Partula t^niata, Morch, is wrongly assigned to the Viti 
Islands. It is peculiar to Moorea, one of the Society Islands. 

Parttjla alabastrina, Pfeiffer, Proc. Zool. Soc. 1S56, p. 39. 
Solomon Isles (Pfeiffer) ; Fiji Islands (Hartmann). 

Partula compressa, Pfeiffer (Bulimus), Zeitschr. f. Malak. 
1850, p. 7.'>. Fiji Islands (Hartmann). 

5. Notes on a small Collection of Shells from the Loo 
Choo Islands. By Edgar A. Smith. 

[Eeoeived February 10, 18S7.] 

Among the valuable collections made at the Loo Choo Islands by 
Mr. H. Pryer were a few shells, which he has liberally presented to 
the British Museum. As five out of the nine species are represented 
by fairly large series of specimens, I have been enabled to make a 
few observations on the variations they present. These may be of 
some use if they tend to prevent the multiplication of species which 
eventually have to be regarded merely as varieties. The specimens 
were obtained, I believe, from the largest island of the group, the 
name of which is variously written Loo Choo, Lu-Tschu, Lu Chu, 
Liew Kiew, and Riu Kin. 

1. Helix despecta, Gray. 

This species, also //. ravida, Benson, H. reJfieldi, H. sieloldiana, 
both of Pfeiffer, and H. ussimilis, H. Adams, are all very much 
alike, and might well be considered varieties of one and the same 



1887.] FROM THE LOO CHOO ISLANDS. 317 

species. In the series of eighteen specimens from Loo Choo I find 
considerable differences in form, some being much higher and more 
globose than others. One example is of a peculiar purplish-brown 
colour, and another is remarkable in having a thickening or limbus 
within the hp. 

2. Helix mercatoria, Gray. 

This species varies considerably in intensity of colour, from a very 
deep black-brown to pale yellowish olive, and the lines of growth in 
some examples are very much coarser than in others. One specimen 
with a comparatively smooth surface has the peripherial dark band 
unbordered by a pale zone on each side as usual. The colour of the 
peristome is also variable, being in the dark or most common forms 
purplish brown, and of a pale flesh-tint in shells of a lighter colour. 

3. Helix luhuana, Sowerby. 

The two specimens which I assign to this species are rather young 
and consequently thinner than adult shells. They differ also from 
the typical form in colouring, having only faint indications of trans- 
verse bauds and much more distinct spiral strise, in which respect 
they exactly resemble H. peliomphala from Japan. 

4. Helix largillierti, Phihppi, var. 

Testa perforata, globoso-conica, mediocriter tenuis, incrementi 
lineis oblique arcuatis tenuiter striata, sordide albida, zonis 
duabus nigro-fuscis cincta, epidermide tenuissima flavescente 
induta ; anfractus 6, convexiusculi, sitperne ad suturam angus- 
tissime submarginati, ultimus subglobosus, ad peripheriam supra 
aperturam obsolete angulatus ; apertura late lunata, longit. 
totius I subcequans ; peristoma expansum, albidum vel dilutissime 
rosea finctum, margine columellari rejlexo, umbilicum setni- 
obtegente. Diam. max. 27 millim., min. 23, alt. 25. 




Helix largillierti, var. 

In form this variety very closely approximates to H. callisona, 
Crosse, but may be slightly higher in the spire. That species, 
however, according to Martens ^ and some specimens from Kiga, 
Japan, which I have assigned to it, is very distinctly spirally striated, 
as is the case in H. peliomphala and other allied species from 

^ Pfeiffer's Novitates Conch, vol. v. p. 31. 
Proc. Zool. Soc— 1887, No. XXII. 22 



318 ON SHELLS FROM THE LOO CHOO ISLANDS. [Mar. 1, 

Japan. On the contrary, the shells here described have a smoother 
aspect and exhibit only the faintest trace of spiral striae. The 
typical form of H. larcjUlierti, also from the Loo Choo Islands, is 
considerably smaller, but of about the same proportions ; it has the 
same number of whorls, a similar umbilicus and angulation at the 
periphery, and the same very faint spiral striation. A variety is 
described by PfeifFer with a single band just above the slight angle 
of the body-whorl, and falling above the sutural line upon the spire. 
The two specimens collected by Mr. Pryer have a similar band, and 
a second situated three or four millimetres below the periphery. 
Although so large, these specimens are evidently young, being very 
thin, and having only partially developed the lip of the aperture ; the 
internal thickening described as present in the type is only feebly 
discernible. 

5. Helix connivens, Pfeiffer. 

Of the forty-six specimens of this species obtained by Mr. Pryer, 
twelve only belong to the unhanded form, the remainder having the 
single red line at the periphery as figured by Reeve and Pfeiffer. 
The lip in both varieties may be either white or pinkish. 

6. Clausilia valida, Pfeiffer. 

Not one of the twenty-four examples of this species at hand 
belongs to the brown-banded variety, all being of a uniform greyish- 
yellow tint. The largest specimen, consisting of seven whorls, is 
33 millimetres in length, or seven longer than the six-whorled 
shell described by Pfeiffer. All have the spire decollated. 

7. Cyclophorus turgidus, Pfeiffer, 

None of the specimens obtained by Mr. Pryer are as large as the 
types, also from Loo Choo, described by Pfeiffer ; but they agree in 
every respect with the small form he mentions from the Ibyat, an 
island of the Bashee group, situated some three hundred miles to 
the south-west of Loo Choo. They appear to be pretty constant in 
form, style of colouring, and in the decided peripherial carination ; 
but the peristome varies from white to a reddish tint, and in some 
specimens it is of a much more duplex character than in others. 
The ojierculum is semitransparent, a little concave externally, and 
consists of seven slowly enlarging whorls which are finely keeled at 
the suture. 

C ibyatensis, Pfeiffer, from the same island as the small variety 
of C. turgidus, differs only in having the last whorl rounded instead 
of more or less carinate at the periphery. It should be regarded as 
a variety of this species rather than as a distinct form. The British 
Museum possesses quite a typical specimen of C. turgidus, from the 
island of Formosa, presented by M. Dickson, Esq. 

8. Cyclophorus exaltatus, Pfeiffer, var. 

A single specimen only was obtained. It differs from Hong-Kong 
and Formosan examples in having a slightly larger aperture, and an 



1887.] MR. H. SAUNDERS ON COSMONETTA HISTRIONICA. 319 

orange-red instead of a white peristome. The convexity of the 
whorls, the sculpture, and the style of colouring are quite of the 
same character. A similar variation in the colour of the peristome 
occurs in the preceding species. 

9. Leptopoma vitreum. Lesson. 

The distribution of this species is very wide, it being recorded 
from the Nicobar Islands, Java, the Moluccas, Philippines, New 
Ireland, Frankland and Fitzroy Islands, and New Caledonia. It has 
not previously been met with at Loo Choo, the most northern point 
of its range yet known. The two specimens are of medium size, and 
of transparent white colour without any markings. 



March 15, 1887. 
Dr. St. George Mivart, F.R.S., Vice-President, in the Chair. 

The Secretary read the following report on the additions to the 
Society's Menagerie during the month of February 1887: — 

The total number of registered additions to the Society's Mena- 
gerie during the month of February was 46, of which 7 were by 
birth, 2 1 by presentation, 7 by purchase, 4 were received on deposit, 
and 7 by exchange. The total number of departures during the 
same period, by death and removals, was 112. 

The most noticeable additions during the month of February were 
as follows : — 

1. A Burmeister's Cariama {Chunga hurmeisteri) , received in 
exchange February 24, being the fifth specimen of this rare species 
of the Northern Provinces of the Argentine Republic that the Society 
has acquired. 

2. A White-fronted Heron {Ardea novcB-hollandicB), from Australia, 
presented by F. B. Dyas, Esq., February 25th. This species is new 
to the Society's Collection. 

3. A young specimen of a Black-winged Kite {Elanus ceeruleus), 
taken from the nest by iMr. R. Southey of Southfield, Plumstead, 
Cape of Good Hope, and received February 28th. This species is 
likewise new to the Society's Collection, 

Mr. Howard Saunders, F.Z.S., exhibited a young male Harlequin 
Duck {Cosmonetta histrionica), shot on the 2nd December, 1886, 
near the Fame Islands, off Northumberland, where it was in company 
with two others, one of which was also obtained (Zool. 1887, p. 70). 
Mr. Saunders stated that records of the occurrence of this species 
were not unfrequent ; but that inasmuch as, with one exception, 
every reputed British specimen which had been submitted to com- 
petent examination had proved to belong to some other species, the 
possessor, Mr. R. W. Chase, of Edgbaston, had kindly complied with 
a request to send his bird up, that its identification might be placed 
beyond a doubt — an example to be commended to other owners of 

22* 



320 MR. o. THOMAS ON BATS [Mar. 15 

rarities. The only British-killed specimen previously known to exist 
was in the collection of Mr. J. Whitaker, of Rainworth Lodge, Notts., 
and bad been obtained at Filey, Yorkshire, in 1862. So-called 
specimens had generally proved to be females or young of the Long- 
tailed Duck or of the American Wood Duck. 



The following papers were read : — 

1. On the Bats collected by Mr. C. M. Woodford in the 
Solomon Islands. By Oldfield Thomas. 

[Received February 11, 1887.] 
(Plates XXV. & XXVL) 

The Mammalian collection made by Mr. C. M. "Woodford in the 
Solomon Islands, and recently acquired by the Natural History 
Museum, consists almost wholly of Bats ; and as nothing has been 
hitherto recorded about the Chiropterous fauna of these islands, his 
collection is naturally of great interest and importance. 

The localities at which Mr. Woodford collected were Alu, in the 
comparatively large Shortland Island, and Faiiro Island, close to 
Shortland, all the specimens therefore coming from the extreme 
western part of the archipelago. 

The collection consists of 23 specimens belonging to 10 species, 
of which two are new, one of these representing also a new genus. 
As might have been expected, the greater proportion of the species, 
and all of those presenting any special interest, belong to the fruit- 
eating section of the order. One Solomon-Island Bat only was not 
obtained by Mr. Woodford, namely Pteropus rai/neri, Gray, which 
comes from the other extremity of the archipelago. This I have 
included in the following list in order to make it a complete catalogue 
of the known species of the group. 

1. Pteropus grandis, sp. n.' (Plate XXV.) 

a, b. Ad. sk,. S and a separate skull. Alu, Shortland Island, 
4/86. 
Size large, about equal to Pt. gouldi. Ears decidedly longer than 
the muzzle, acutely pointed. Origins of wings about an inch apart 
n\ the back. Interfemoral membrane very narrow in the centre, 
concealed by the fur. Fur rather coarse, hispid over the shoulder- 
glands, rather woolly on the legs. Fore limbs and membranes 
nearly naked above, a few scattered black hairs on the proximal 
half of the forearm. Fur on back adpressed, rather more than an 
inch in breadth at its narrowest part. Rump and hind limbs nearly 
to the ankles thickly clothed with woolly hairs. Below, the 
humerus, proximal half of forearm, and the membranes between the 
humerus and femora are covered with hair, and a thin baud of fur 

1 Preliminary diagnosis published, Ann. & Mag. N. H. (5) sis. p. 147, Feb. 

1887. 



T.Z.S.1887. PI. XXV 




J.SmiL liLh 



PTEROPUS GKAl^TDIS. 



Haniharfc imp 






I* T^ 



cO/ 



P,Z.S.1887.P1.XXVI 



\ 



s. 




'^/ 






JSmiLliL}. 



NESONYCTERTS WOGDFORDJ 



ffirihart imo. 



1887.] 



FROM THE SOLOMON ISLANDS, 



321 



passes along outside the forearm nearly to the wrist. Femora and 
proximal third of tibiee thickly clothed with softer woollier fur. 
■ Face, middle of back, chin, and centre Hue of belly black. Neck 
all round, shoulders, sides of body, and hind limbs below dark ma- 
roon-red, the tufts over the shoulder-glands rather paler. Rump 
and upperside of thighs bright orange-yellow, coutrasliug markedly 
with the sombre hue of the back. 

Skull agreeing closely with that of ft. goiildi in size and pro- 
portions. 

Teeth (fig. 1) large and heavy as compared to those of Ft. gouldi, 
Pt. chri/soproctus, and others, but far smaller and lighter than those 
of PL vielanopogon. Upper incisors broad, touching each other, the 
combined breadth of the four 8 mm. Canines rather short, thick 
and stout, with a broad and prominent internal basal ledge. Anterior 
premolars minute, deciduous. Second premolar large and broad, 
with a prominent internal secondary cusp. All the cheek-teeth 

Fig. 1. 





Teeth of Fteropus grandii, natural size. 

broad and strong ; combined length of the three largest 18 mm., and 
the breadth of the centre one 4 mm. Last molar slightly larger 
than one of the outer incisors, its antero-posterior diameter 2*2 mm. 
Lower teeth presenting very much the same characters as the upper. 
Outer incisors very large, 2 mm. in transverse diameter. Canine 
with large internal basal ledge or cingulum as in the upper jaw. 
Second premolar unusually near the canine, the comparatively large 
first premolar larger than the diastemata in front of and behind it, 
2*6 mm. in diameter. Last molar about equal to one of the middle 
upper incisors, smaller than the outer and larger than the inner 
lower incisors. 

Dimensions of an adult male in skin : — Head and body (c.) 325 
mm., head 74, muzzle 32, ear (above crown) 30, forearm 170 
( = 6*7 in.), thumb 73, index finger 118, tibia 122, calcaneum 24. 

Skull (specimen b). Basal length 67, greatest breadth 39, supra- 
orbital foramen to tip of nasals 30, interorbital breadth 10, inter- 
temporal breadth 7"5, breadth from tip to tip of postorbital processes 
30, palate length 40. 



322 MR. O.THOMAS ON BATS [Mar. 15, 

It is with considerable reluctance that I find myself compelled to 
add another species of Pteropus to the long list of those already 
known ; but the characters of Pt. grandis so entirely fail to fit in with 
those of any of the hitherto described species, that I have no alter- 
native but to do so. 

Pt. (jravdis differs from every known species at all approaching 
its size by its dark maroon-coloured neck, throat, and sides, and by 
its bright yellow rump. Apart from coloration, again, it differs 
from Pt. edulis by its much smaller size and broadly edged canines, 
from Pt.gouldi, aneiteanus, and poliocephalus by its very much heavier 
teeth, and from Pt. inelanopogon by its smaller teeth and longer 
pointed ears. On the whole it may be looked upon as most nearly 
allied to Pt. chrysopi-octus, a native of the Moluccas, which resembles 
it in many of its characters, but differs by having its neck both above 
and below rich yellow, by its yellowish crown and dark-coloured 
rump. The teeth also of Pt. chrysoprocius are smaller and lighter 
than in Pt. grandis ; tlie canines are thinner and have narrow 
postero-internal ledges, and, finally, there is a much greater space 
between the canines and second premolar below, the anterior pre- 
molar having a diameter less than the length of either of the diaste- 
mata in front of or behind it. 

To another species also Pt. grandis bears a certain amount of 
resemblance, namely to Pt. rai/neri. Gray, also from the Solomon 
Islands ; but that species has much shorter ears, and is very far 
smaller, having a forearm only 135 mm. long, a skull only 55 mm. 
long, and teeth which, although they have very much the same 
shape and relative proportions as in Pt. grandis, yet differ so 
markedly in their actual size as to preclude all possibihty of the two 
species being the same. 

2. Pteropus hypomelanus, Temm. 
a. Alu, Shortland Island, 4/86. 

Previously known range, from Borneo to New Guinea. 

This is the first published notice of the occurrence of this species 
in the Solomon Isles ; but its discovery there was made in 1883, when 
Surgeon H. B. Guppy, of H.M.S. ' Lark,' obtained and sent to the 
Museum a specimen, also collected on Shortland Island. 

3. Pteropus rayneri, Gray^. 

Discovered in the islands of San Christoval and Guadalcanar in 

^ I may take this opportunity of stating that an examination of the typical 
specimen of Pteropus molosdmis, Temm., preserved in the Leyden Museum, 
proves that the Caroline-Island Pfcro]7V.s described by me in 1S82 (P. Z. S. ' 
1882, p. 756) under the name of Pt. hrevicepg is not really distinguishable from 
that species, of which, tip to that date, the locality was unknown. I must, 
however, for my own justification, point out that the shoulder-tufts of Pt. mo- 
lossinus, instead of being " bright yellow" as has been described, are really of a 
dark orange-brown, but little in the type, and in my specimens not at all, 
ligliter tliau the general colour of the body. Nor can I at all fully appreciate 
the alleged resemblance in dentition between the very small-toothed Pf. molos- 
sinus (see figures t.c. pi. Iv.) and the large-toothed Pt. aneiteanus and Pt.Ju- 
baiws, the latter of which has the largest teeth of any member of the genus. 



18S7.] FROM THE SOLOMON ISLANDS. 323 

1854 by Dr. F. M. RHvner, during the voyage of H.M.S. ' Herald ;' 
not obtained by Mr. Woodford. 

4. Cynonycteris brachyotis, Dobs. 
a, h. Fauro Island, 5/86. 

Preriously known habitat, New-Ireland group ; also recorded from 
Celebes ^ 

The two specimens obtained by Mr. Woodford are both slightly 
immature, and both retain their first upper premolars. They agree 
in every respect with the typical specimens. 

5. Harpyia major. Dobs. 

a, h. Alu, Shortland Island, 4/86. (New Georgia, Coll. Brit. 
Mus.) 
Hah. E. New Guinea ; New-Ireland group ; Solomon Islands. 

6. Cephalotes peronii, Geoffr. 

a. Alu, Shortland Island, 4/86. (Ugi, Dr. H. B. Guppy, 
H.:SI.S. 'Lark'; San Christoval, F. M. Rayner, H.M.S. 
'Herald.') 
Hah. Austro-Malayan subregion, from Celebes to Solomon 
Islands. 

Nesonycteris", g. n. 

Muzzle long, narrow, cylindrical ; nostrils projecting considerably ; 
upper lip with a vertical groove bounded laterally by raised naked 
edges ; posterior palate-ridges divided in the centre ; index finger 
witbout a claw, longer than the metacarpal bone of the middle finger ; 
wing-membranes as in Melonycteris ^ ; tail none. 

Dentition :— I. f, C.\, Pm. |, M. |x2=32. 

Shape, size, and position of teeth much as m llelonycteris^, but 
lower inner incisors entirely obsolete. 

Skull as in Melonycteris; premaxillfe distinctly separated ante- 
riorly \ Lower jaw with a very long gutter-like symphysis, and with a 
long diastema between the first premolar and that next behind it. 

This most interesting new genus presents a combination of the 
characters of several of the hitherto known Macroglossine Bats. 
Thus, in the clawless state of the index finger it resembles Eotiyc- 
teris and Notopferis ; by the attachment of its wing-membranes and 
the form of its palate-ridges, Melonycteris and, fairly closely, Mega- 
Inglossus. And, again, while the number of the incisors is as in 
Notopteris, the other teeth closely resemble those of Melonycteris, 
to which, on the whole, it is certainly most nearly allied. 

1 Jentink, Notes Leyd. Mus. t. p. 173 (1883). 

^ viiaos, an island, vvKTepb, a bat. 

3 See Dobson, P. Z. S. 1877, p. 119. 

■* Figured, id. I. c. figs. 5 and 6. 

^ As tbey are in Melonycteris, altbougb by some accident that genus lias been 
placed under the heading of " premaxillary bones united in front," in the Cat. 
Chiropt. Brit. Mus. p. 4, nothing being said on the subject in the description 
of the genus pp. 97-8. 



324 MR. o. THOMAS ON BATS [Mar. 15, 

This discovery and that of the highly interesting West-African 
Megaloglossus ivoermanni, Pageust.\ of which I have seen a beauti- 
ful spirit specimen from Liberia in the Leyden Museum, render the 
synopsis of genera in Dr. Dobson's Catalogue of Chiroptera somewhat 
obsolete, since, according to Dr. Dobson's synopsis, Nesomjeteris would 
eome next to Notopteris and I^onycteris instead of to Melonycteris, 
its nearest ally. I have therefore thought it convenient to draw up 
the following synopses of the Macroglossine genera, the first based 
solely on the soft-part and external characters, and the second on 
those of the skull and teeth. 

I. External or Soft-part Characters. 

A. Tail very sbort ; wings from the sides of the back. 
a. Wing-membrane from the base of the first toe ; 

no claw on index 1. EoNYCTERrs, 

h. Wing-membrane from the third, or second and 
third toes. 
a'. A claw on index. 

a". Posterior palate-ridges undivided 2. Macroglossus, 

h". Posterior palate-ridges divided in centre. 
a'". Lower jjart of rhinariiim broad, convex 

laterally 3. Megaloglossus. 

h'". Lower part of rhinarium narrow, concave 

laterally 4. Melonycteris. 

h' . No claw on index. Palate-ridges and rhinarium 

as Melonycteris .5. Nesonycteeis. 

B. Tail long ; wings from centre of back ; no claw on 

index 6. Notopteris. 

II. Cranial and Dental Characters. 

A. Pm.^ above and below small, single-rooted. 

a. Penultimate molar many times as large as last 1. Eonycteris. 

b. Penultimate molar but little larger than last. 

a'. Vm.} nearly as large as that next behind it ; 

premaxilla; united 2. MacroglossuSv 

b'. Vn\} minute. 

a" . Middle premolar three fourths as long as the 

canine 3. Megaloglossus^. 

b". Middle premolar barely one third as long as 
the canine ; premaxillaj separate . 

a'". Incisors I 4. Melonycteris. 

h'". Incisors Y 5. Nesonyctbris. 

B. Pm.i above and below long, double-rooted 6. Notopteris. 

7. Nesonycteris woodfordi, sp. n.' (Plate XXVI.) 

a. Ad. sk. $ . Fauro Island, 5/86. 

b, c. Ad. sk. d" and yg. al. Alu, Shortland Island, 4/86. 
Strikingly like Melonycteris melanops\ Dobs., in size, proportions, 

shape and length of ears, and in the colour and texture of the fur of 

' JB. Hamb. ii. p. 125, pi. i. (1885). 

"^ For several details connected with the dentition of this interesting genus, 
I am indebted to Dr. F. A. Jentink, of the Leyden Museum. 

^ Preliminary diagnosis jmblished. Aim. & Mag. N. H. (5) xix. p. 147, Feb. 1887. 
^ Figured, P". Z. S. 1877, pi. xvii. 



1887.] 



FROM THE SOLOMON ISLANDS. 



325 



the back, this being of precisely the same soft cottony nature, and 
of the same fulvous-yellow tinge ; but the face, instead of being 
yarigated with black and white, and quite different from the back, 
is quite like the latter, except that it is rather darker. The under- 



Fis. 2. 




Skull of Kesonj/cfcris woodfordi, upper view ; twice natural size. 



Fig. 3. 




Skull of Nescmycteris woodfordi, side view ; twice natural size. 



side also is very similar to the back, although rather paler and 
duller, while in M. tnelanops it is nearly black. Point of insertion 
of the antebrachial membrane without any trace of a white spot. 



326 MR. O.THOMAS ON BATS [Mar. l.*). 

Ears small, obtusely pointed. Nostrils very prominent. Palate- 
ridges 7 in number, the 6th and 7th divided in the centre as in 
ilf. melanops^. 

Humerus, proximal half of forearm, and upperside of hind legs 
to ankles thickly clothed with soft woolly fulvous fur. Wing- 
membranes behind humerus and whole of interfemoral membrane 
also covered with fur. On the lower side the same parts are hairy 
as on the upper, but the hair is much thinner and sparser. 

Skull (figs. 2 and 3) long and slender, with a long narrow muzzle ; 
postorbital processes well developed ; premaxillse widely separated in 
front. 

Upper incisors 4, forming an even semicircular row ; canines very 
long and powerful, with deep vertical grooves on their anterior, 
externa], and posterior surfaces ; pm.^ minute, quite close to the 
canine ; molars verj' small, smaller even than in Melonycteris, 
posterior one about two thirds the size of the anterior. 

Lower incisors one on each side, near the canines, widely separate 
in the middle ; canines slightly grooved posteriorly ; pm.' close to 
the canines, and succeeded by a broad diastema ; posterior molar in 
section of about the size of pm.\ 

Dimensions of specimen b, an adult male (skin) : — 

Head and body (c.) 100 mm., head 35, muzzle 15, ear (from notch 
at base) 11, above crown 8, forearm 55 ( = 2*18 in.). 

8. Phyllokhina diadema, GeofTr. 

a. Fauro Island, .5/86. 
Previously known range : Oriental Region as far east as the Key 
Islands and Western New Guinea. 

9. Phyllorhina cervina, Gould. 

a-d. 3 6 & 1 2 . Fauro Island, 5/86. 

Previously known range : N. Australia, New Guinea and neigh- 
bouring islands, including Duke of York Island. 

One of the male specimens has no transverse frontal sac, while the 
other two have it well developed. In all other respects, however, 
the specimens are quite identical. 

10. Vesperugo abramus, Temm. 

a. Fauro Island, 5/86. 
Previously known range : Palsearctic, Oriental, and Austro-Malayan 
part of Australian Region, as far east as New Guinea. 

11. Emballonura nigrescens, Gray. 

a-f. All $. Fauro, Shortland, and Savo Islands. (Ugi; 
Guppy.) 
Previously known range : Austro-Malayan subregion, as far east as 
New Ireland. 

This species seems to be very common in the group, as both 

^ See figure by Pagenstecher, Naturh. Mus. Hamb. 1884, tab. fig. 2. 



1887.] FROM THE SOLOMON ISLANDS. 327 

Surgeon Guppy and Mr. Woodford obtained it in considerable 
numbers. 

For a comparison of the Chiropterous fauna of the Solomons with 
that of the neighbouring islands, it fortunately happens that the 
Bats of the nearest group, viz. that of New Britain, New Ireland, and 
Duke of York, have been fully worked out by Dr. G. E. Dobson^, 
who based his papers on the specimens obtained in those islands by 
the Rev. George Brown. These specimens are all in the Natural 
History Museum, so that I have had the advantage of being able to 
compare Mr. Woodford's Bats directly with those named by the 
chief living authority upon Chiroptera. 

The following parallel lists show the species as yet known from 
the two groups, those marked with an asterisk being peculiar to 
their respective groups. 

Megachiroptera. 

New-Ireland group. Solomon group. 

Pteropus melanopogon. *Pteropvis grandis. 

capistratus. hypouielanus. 

* rayneri. 

CjTionycteris brachyotis. Cynonycteris bracbyotis. 

Harpyia major. Harpyia major. 

Cephalotes peronii. Ceplialotes peronii. 

Macroglossus minimus. 
*Melonjcteris melanops. *N'esonycteris woodfordi. 

MiCROCHIROPTERA. 

Phyllorbina trieuspidata. Phyllorhina diadema. 

cervina. — — cervina. 

calcarata. 

*Vesperugo angulatus^. Vesperugo abramus. 
Kerivoula bardwickii. 

Emballonura nigresoens. Emballonura nigrescena. 

The New-Ireland group has therefore two, and the Solomon group 
three peculiar species, while there are five species common to both 
groups, a number that is certain to be largely increased as the 
islands are more fully explored. The proportion of fruit-eating to 
insectivorous Bats is larger by a slight fraction in the Solomons than 
in the other group, a difference only to be expected from the more 
oceanic position of the former. This position has also resulted, so 
far as is yet known, in the nearly entire absence of terrestrial Mam- 
malia in the Solomons, the only other mammals besides Bats known 
from there being the arboreal and widely-spread discus orientalis, 
Pall., and a Rat from Florida Island, described by Mr. E. P. Ram- 
say'. On the other hand, Mr. Brown collected in the New-Ireland 
group, as recorded by Mr. Alston*, no less than six indigenous 

1 P. Z. S. 1877, p. 114, and 1878, p. 314. 

^ Peters, SB. nat. Freund. 1880, p. 122. Only known to me by tbe original 
description. 

3 Proo. Linn. Soc. N. S. W. vii. p. 43, 1882. This Eat appears to be a 
member of tbe arboreal genus Uromys. 

* P. Z. S. 1877, p. 123. 



328 MR. W. R. OGILVIE-GRANT ON BIRDS [Mar. 15, 

species of Rodents and Marsupials, many of them of such a distinctly 
non-arboreal nature as to preclude the possibility of their having 
been originally introduced on drifting logs or trees, a means of dis- 
tribution to which no doubt the Solomons owe the presence of their 
two non-flying mammals and Rat, the Cuscus. 



2. A List of the Birds collected by Mr. Charles Morris 
Woodford in the Solomon Archipelago. By W. R. 
Ogilvie-Qeant, 

[Eeceived February 14, 1887.] 
(Plate XXVII.) 

The Natural History Museum has recently received a collection 
of Birds made by Mr. C. M. Woodford at Fauro, Alu, Shortland 
Island, and other localities ; and, although comparatively few 
specimens were obtained, some of the species are very interesting, and 
one at least is new to science. This is a Crow belonging to the 
genus Macrocoi-ax. 

The following is an extract from a letter received from Mr. C. M. 
Woodford :^ — "I find that Hornbills and Cockatoos do not extend 
beyond Malayta, being entirely unknown on the island of San Chris- 
toval and the smaller islands adjacent ; and as they are not found 
in the groups to the south-east, this will be the limit to which these 
two genera extend." 

The following is a list of the species, with remarks on some of the 
rarer ones, and notes on their soft parts made by the collector. 

1. ASTUR PULCHELLUS. 

Urospizias pulchellus, Ramsay; Salvad. Orn. Papuasia, iii. App. 
p. 508. 

a. S ad. Alu. Iris brown ; bill black at tip ; nostrils and base 
of mandibles yellow ; legs yellow. 

b. (S juv. Fauro. Iris brown ; legs and base of bill yellow ; the 
tip black. 

This species, at first described by Ramsay as A. soloensis. Lath., 
as already observed by Finsch, is a very distinct bird. It is nearly 
allied to 'A. dampieri, Gurney, but may be easily distinguished by 
the colour of the breast and abdomen, which is a uniform deep 
vinous red (chestnut-hazel, Ridgway) instead of vinous with faint, 
narrow, pale cross bars on the flanks, belly, and under tail-coverts. 

Char. Male adult. Head, back, upper coverts, quills and tail- 
feathers slaty grey (Ridgway), the latter without any trace of cross 
bars. Throat and fore neck light slaty grey. Breast, abdomen, 
flanks, and under tail-coverts deep vinous red (chestnut-hazel, 
Ridgw.). Axillaries and under wing-coverts pale vinous red the 
latter with greyish cross bars. Quills below slaty grey, the inner 
web lighter, with whitish wavy bars towards the distal extremity. 

Total length 13-5 inches, culmen -6, wing 77, tail 6*25, tarsus 2-1. 



p. z.s;i887. pi.ixvn. 




^^' 



*HBt ad rt, Kih 



MACROCORAX WOODFORD! 



Minterrv Broa , imp. 



1887.] FROM THE SOLOMON ARCHIPELAGO. 329 

Young male. Greneral colour above warm brown, with rufous 
edges to the feathers. Upper wing- and tail-coverts brown, with 
rufous edges. Quills above brown, with narrow light-red edges, 
indistinctly barred with dark brown, except on the outer web of the 
primaries, the distal half of the inner web pale huffy red, showing 
the dark bars very strongly. Tail-feathers brown, with 9 or more 
narrow dark brown cross bars, the outer pair on each side inclining 
to reddish. Crown of head, occiput, and sides of neck dull brown. 
Nape and hind neck reddish brown, the former showing traces of 
grey. Lores, eyebrows, ear-coverts, and cheeks grey. Throat 
whitish grey. Neck, breast, and under tail-coverts huffy white, with 
broad reddish-brown bars on some of the feathers. Abdomen 
whitish buff and almost immaculate. Sides of body, flanks, and 
thighs reddish buff with faint bars of darker. Under wing-coverts 
and asillaries the same. Under surface of quills ashy white, shading 
into pale huffy red, with marked brown cross bars, indistinct on the 
outer web of the primaries. 

Measurements the same as above. 

This young male is evidently just entering its first moult, and has 
already got one of the adult grey feathers in the secondaries. 

2. Haliastur girrenera. 

Haliastur girrenera (V.) ; Salvad. t. c. i. p. 15. 

a, b. 5 ad. Alu. Iris brown ; bill and legs yellow. 

c. c? juv. Fauro. Bill black ; iris brown ; legs yellow. 

3. Pandion leucocephalus. 

Pandion leucocephalus, Gould; Salvad. t.c. i. p. 11. 
a, b. S Alu. Iris pale or dull yellow ; bill black ; legs dirty 
grey. _ . _ _ 

c. d" juv. Alu. Iris reddish yellow. 

4. Cacatua ducorpsi. 

Cacatua ducorpsii, Jacq. & Pucher. ; Salvad. t. c. i. p. 104. 
a. c? ad. Alu. Iris brown ; skin round eye blue ; bill and feet 
grey. 

5. Geoffroyixjs heteroclitus. 

Geoffroyius heteroclitus (Hombr. & Jacq.) ; Salvad. t. c. i. 
p. 194. 

a. c? ad. Alu. 

b, c. 2 ad. Alu. Iris pale yellow ; bill black ; feet and legs 
grey. 

6. ECLECTUS POLYCHLORUS. 

Eclectus pectoralis (P. L. S. Miill.) ; Salvad. t. c. i. p. 197- 

a. c? ad. Alu. Upper mandible yellow ; legs black ; iris yellow. 

b, c. S ad. Alu. Upper mandible orange, lower black ; legs 
dirty grey; iris yellow. Norn, vernac. " Karo." 



330 MR. W. R. OGILVIE- GRANT ON BIRDS [Mar. 15, 

d. 2 ad. Alu. Bill and feet black ; iris yellow. 

e. 2 ad. Shortland Island. Legs and bill black ; iris yellow. 

f. (5 ad. Fauro. Iris yellow ; upper and lower mandibles 
yellow ; feet black. 

7. Eos CARDINALIS. 

Uos cardinalis (Gray) ; Salvad. t. c. i. p. 249. 
a, h. (S and $ ad. Iris red ; feet black ; bill yellow and black. 
c. 5 ad. Alu. Iris red ; legs black ; bill black with yellow 
tip. 

8. Trichoglossus massena. 
Trichofflossus massena, Bp. ; Salvad. t. c. i. p. 288. 

a, b. 6 ad.; c. ? ad. Alu. Iris red ; bill orange; legs grey. 

9. Rhytidoceros plicatus. 

Rhytidoceros pUcatus (Penn.); Salvad. t. c. i. p. 392. 

a. (S ad. Shortland Island. Iris yellow; legs black; bill dirty 
white, reddish at base ; skin white at eyes and throat. 

b. 2 ju"^- Gruadalcanar. 

10. Halcyon satjrophaga. 

Sauropaiis saurophaga (Gould) ; Salvad. t. c. i. p. 469. 
a, b. 6,2 ad. Fauro. Iris brown ; legs and bill black. 
The female is very distinctly greener on the back than the male, 
which is much brighter blue. 

11. Halcyon TRiSTRAMi. 

Sauropaiis tristrami (Layard) ; Salvad. t. c. iii. App. p. 524. 
a, b. S ,2 ad. Fauro. Iris brown ; bill and feet black ; the 
lower mandible whitish at base. 

12. Halcyon sancta. 

Sauropaiis sancta (Bodd.) ; Salvad. t. c. i. p. 470. 

a. 2 ■ -Alu- l^'"is ''ark brown ; feet and bill black above, the 
latter white below. 

b, c. S • Fauro. Iris brown ; bill black, whitish below ; legs 
black. 

13. EURYSTOMUS crassirostris. 

Eurystomus crassirostris, Sclater ; Salvad. t. c. i. p. 510. 

a, b. 6 ad. ; c, d. 2 ad. Alu. Iris brown ; bill and feet red. 

14. Macropteryx mystacea. 

Macropteryx mystacea (Less.); Salvad. t. c. i. p. 537. 
a. 2 ad. Alu. Eyes black ; bill and feet black. 

15. HiRTJNDO TAHITICA. 

Hirundo taMtica, Gm. ; Salvad. t. e. ii. p. 5. 
a. 2 ad. Alu. Bill, legs, and iris black. 



J887.] FROM THE SOLOMON ARCHIPELAGO. 331 

16. MONARCHA CASTANEIVENTRIS. 

Pomarea castaneiventris (Verr.) ; Salvad. t. c. ii. p. 11. 

a. 2 ad. Alu. Iris brown ; bill aud legs bluish grey. 

b. 5 ad. Fauro. Iris brown ; bill grey ; legs black. 

This specimen is referred with doubt to the above-named species, 
being a somewhat smaller bird, and having the chestnut of the breast 
and belly much darker than the type. 

I 7. RhIPIDURA TRICOLOR. 

Saidoprocta melaleuca (Q. & G.) ; Salvad. t. c. ii. p. 48. 

a. 2 ad. Uru Bay, Malayta. Legs, beak, and iris black. 

With nest, taken from the upper part of a dead limb of a tree, 
which it assimilated in colour, and two eggs, partially incubated, 
one of which was broken. 

18. Rhipidura rtjssata. 

Rhipidura russata, Tiistr. ; Salvad, t. c. ii. p. 67. 
a. Shortland Island. Iris black. 

19. Graucaltjs pusilltjs. 

Graucalus pusillus, Ramsay; Salvad. t. c. ii. p. 140. 

a. S juv. Alu. Legs and bill black ; iris pale yellow. Nom. 
vernac. " Binbin." 

b, c. (S and 5 • Fauro. Bill and feet black, iris yellow. 

20. Graucalus elegans. 

Graucalus elegans, Ramsay, Proc. Linn. Soc. N. S. W. vi. p. 176. 
a. (S ad. Fauro. Iris brown ; legs and bill black. 
Agrees well with Mr. Ramsay's description. 

21. Calornts metallica. 

Calornis metallica (Temm.) ; Salvad. t. c. ii. p. 447- 

a. 2 ad. Alu. Bill and legs black ; iris bright red. Nom. 
vernac. " Ouriiiri." 

b, c. rS , 6 juv. Fauro. Iris red; bill and legs black. 

22. Calornis cantoroides. 

Calornis cantoroides, G. R. Gr. ; Salvad. t. c. ii. p. 456. 

a. 5 ad. Alu. Bill aud feet black ; iris red. 

I have compared the above specimen, which is evidently C. 
solomonensis of Ramsay, with Gray's types of cantoroides in the 
British Museum, and find them in every respect similar. 

23. Lamprocorax grandis. 

Lamprocorax grandis, Salvad.; Salvad. t. c. ii. p. 460. 

a. Alu. Iris red ; bill and legs black. 

b, c. (S , 2 ad- Fauro. Iris dark red ; bill and feet black. 



332 ON BIRDS FROM THE SOLOMON ARCHIPELAGO. [Mat. 15, 

24. MiNO KREFFTI. 

Mino kreffli (Sclat.) ; Salvad. t. c. ii. p. 469. 

a. $ ad. Alu. Legs, beak, and patch round the eye bright 
yellow ; iris yellow, pupil black. Nom. vernac. " Tigeno." 

b. S ad. Fauro. Iris, legs, and bill bright yellow. 

c. 2 ad. Fauro. Iris, eye-patches, legs, and bill bright orange. 

25. Macrocorax woodfordi, sp. nov. (Plate XXVII.) 

a. 2 ad. Aola, Guadalcanar. Iris dark grey ; bill grey (yellowish 
in the skin, with black tip) ; legs black. 

The general colour above is black glossed with greenish on the 
head and neck and purplish blue over the rest of the upper surface. 
Head, neck all round, underparts, flanks, thighs, and under tail- 
coverts black, glossed with green. The rest of the feathers black, 
glossed with purplish blue, except some of the feathers of the wing- 
coverts, back, secondaries, and primaries, which are dull brownish 
black and evidently worn, as the bird is in full moult. 

Total length 15| inches, culmen 2-3, wing 10-3, tail 5, tarsus 2. 

This species is very strongly marked, being one third smaller than 
the only other known species 31. fuscicapUlus, Gray, from the Aru 
Islands, from which it also differs in general colour and in having 
the upper mandible less strongly arched. 

26. Ptilopus lewisi. 

Ptilopus lewisii, Ramsay ; Sharpe in Gould's ' Birds of New 
Guinea,' pt. xvii. 

a, b. c? ad. Alu. Bill yellow ; feet red ; iris yellow. 

27- Globicera rufigxjla. 

Carpophaga rufgula, Salvad. t. c. iii. p. 79. 

a. c? ad. Malayta. Bill, wattle, legs, and iris red. 

This specimen agrees well in every respect with the type specimen 
in the British Museum, and is easily distinguished from the nearly 
allied G. rubricera of G. R. Gray, from New Ireland, by the burnished 
green-gold back, pale vinous throat, and dove-grey chest, while in 
the latter the back is burnished red-gold and the throat uniform pale 
vinous with the chest. 

28. Macropygia rufo-castanea. 

Macropygia rufo-castanea, Ramsay; Salvad. t. c. iii. p. 149. 
a. S ad. Alu. Eye brown; bill and legs brown. 

29. Calcenas nicobarica. 

Calcenas nicobarica (Linn.) ; Salvad. t. c. iii. p. 209. 
a. d" ad. Alu. Iris pale yellow ; bill and feet black. 

30. Megapodius brenchleyi. 

Megapodius brenchleyi, G. R. Gr. ; Salvad. t. c. iii. p. 240. 
a. 2 ad. Alu. Bill dull yellow ; legs black ; iris brown. 



p. Z.S.1887. P]. XT/Ill 




Peter Snot del. et htt 

1. LEPID0DACTY1.US WOODFORDll. 2 . TYPHLOPS ALUENSIS 
3.BATF1ACHYL0DES VERTEBRALIS. 4. ITfLA LUTEA, 



Mintern Bros , in^ . 



1887.] ON THE HERPETOLOGY OF THE SOLOMON ISLANDS. 333 

31. PORPHYRIO MELANOPTERUS. 

Porphtjrio melanopterus, Temm. ; Salvad. t. c. iii. p. 280. 
a. cS arl. Fauro. Iris reJdish brown ; bill and shield red ; legs 
dirty pink ; joints of legs and toes grey. 

32. Demiegretta sacra. 

Demiegretfa sacra (Grmel.) ; Salvad. t. c. iii. p. 345. 

a. (S ad. Fauro. Iris yellow ; legs yellow ; bill yellowish black. 

33. BUTORIDES JAVANICA. 

Butorides javanica (Horsf.); Salrad. f. c. iii. p. 359. 
a. tS ad. Fauro. Upper mandible black, lower one pale yellow ; 
iris yellow ; feet yellow ; legs grey. 

34. Nycticorax caledonicus. 

Nycticorax caledonicus (Gmel.); Salvad. t. c. iii. p. 3/2. 

a. (S juT. Fauro. Iris yellow ; legs yellow ; bill black, yellow 
below. 

35. Angus leucocapillus. 

Anous leucocapillus, Gould ; Salvad. t. c. iii. p. 457. 
a. Shot at sea. Bill, eyes, and legs black. 



3. Second Contribution to the Herpetology of the Solomon 
Islands*. By G. A. Boulenger, F.Z.S. 

(Plate XXVIJI.) 

[Eeceived February 15, 1887.] 

Eicli as has been the berpetological booty of Mr. H. B. Guppy's 
exploration of the islands of Bougainville Straits, the knowledge of 
that fauna is so far from being exhausted that a recent visit to the 
same islands by a second collector, Mr. C. M. Woodford, has yielded 
examples of as many as nine more species of Reptiles and Batrachians, 
seven of which are altogether new to science. The collection now 
reported upon was made at two localities, viz.. Faro Island, and Alu, 
Shortland Islands. It contains specimens of the following species 
already known from the Solomon Group : — 

Faro: — Gymnodactijlus pelagicus, Gir., Gehyra oceanica. Less., 
Lepidodactylus guppyi, Blgr.^ Geclco vittatus, Houtt., Varaiius 
indicus, Daud., Lygosoma smaragdinum. Less., L. cyaiiurum. Less., 
L. cyanogaster, Less, {carteretii, D. & B.), L. nigrum, Hombr. & 
Jacq., Dipsas irregularis, Merr., Hoplocephalus j^ar, BIgr., Rana 

» Cf. Tr. Zool. Soc. xii. pp. 35-62, pis. vii.-xiii. (1886). 

^ Male with a long, uuinterrupted series of femoral and pra\inal pores, 
angulur mesially ; 44 pores aUogftlier. 

Proc. Zool. Soc— 1887, No. XXIII. 23 



334 MR. G. A. BOULENGER ON THE [Mar. 1.5, 

hufoniformis, Blgr., U. gvppyi, Blgr.\ Cornvfer giippyi, Blgr., 
C. sofotnonis, Blgr., Ceratobutrachus guentheri, Blgr. 

AIu : — Corucia zehrata. Gray, Dendrophis solomonis, Gtlir., 
Dipsas irregularis, Merr. 

In addition to these species, the following, new to the Solomons, 
■were obtained : — 

REPTILIA. 

Lepidodactylits woodfordi, sp. n. (Plate XXTIII. fig. 1 .) 

Closely allied to L. gvppyi. Digits without distinct web. Tail a 
little depressed, rounded. Femoral and praeanal pores 25 altogether. 
Grey above, with zigzag blark cross bands, six between the nape and 
the base of the tail ; a black streak from the nostril to the neck, 
passing through the eye and above the ear ; lower surfaces whitish. 

millim. 

Total length 78 

Head 11 

AVidtli of head 7 

Body 29 

Fore limb 14 

Hind limb ,. 18 

Tail 38 

Faro Island. A single male specimen. 

Lygosoma solomonis, sp. n. 

Body elongate, limbs short ; the distance between the end of the 
snout and the fore limb is contained once and three fifths to once 
and two thirds in the distance between axilla and groin. Snout sliort, 
obtuse. Lower eyelid scaly. Nostril pierced in a single nasal ; no 
supranasal ; a single anterior loreal ; frontonasal broader than long, 
forming a narrow suture with the rostral and with the frontal ; latter 
shield about as long as frontoparietals and interparietal together, in 
contact with tlie first and second supraoculars ; four supraoculars ; 
seven snpraciliaries ; frontoparietals and interparietal distinct, latter 
a little smaller than former; parietals forming a suture behind the 
interparietal ; four to six pairs of nuchals ; fourth or fifth labial 
below the eye and entering the orbit. Ear-opening oval, a little 
smaller than the eye-opening ; no auricular lobules. 24 or 26 
smooth scales round the middle of the body, the two vertebral series 
largest. A pair of large praeanals. Limbs widely separated when 
adpressed ; tiie length of the hind limb equals the distance between 
the anterior border of the orbit and the fore limb. Digits short; 
subdigital lamellpe smooth, undivided, 15 to 17 under the fourth toe. 
Tail thick, once and one third the length of head and body. Brown 
above, dotted with blackish ; pale brownish inferiorly, dotted with 
brown. 

^ One of the specimens exceeds tbe type {I. c. pi. ix.) in size, measuring 185 
millim. from snout to vent. 



1887.] HERPETOLOGY OF THE SOLOMON ISLANDS. 335 

niillim. 

Total length 135 

Head 11-5 

Width of head lf> 

Body 45-5 

Fore limb 9 

Hind limb 16 

Tail 78 

Three specimens from Faro Island. 

Lygosoma woodfordi, sp. n. 

Body elongate, limbs rather short ; the distance between the end 
of the snout and the fore limb is contained once and three fifths in 
the distance between .ixilla and groin. Snout moderately elongate, 
truncate. Lower eyelid scaly. Nostril pierced in a single nasal ; 
no supranasal ; a single anterior loreal ; rostral forming a broad 
straight suture with the frontonasal, which is broader than long ; 
prsefrontals forming a short median suture; frontal as long as fronto- 
parietals and interparietal together, in contact with the first supra- 
ciliary and the two anterior supraoculars ; four supraoculars, followed 
by a very small fifth, first longest ; ten supraciliaries, first largest ; 
frontoparietals and interparietal distinct, former much larger than 
latter ; parietals forming a suture behind the interparietal ; no 
nuchals ; nine upper labials, seventh below the centre of the eye ; 
a series of rather large suborbitals separates the orbit from the 
labials. Ear-opening oval, a little smaller than the eye-opening ; no 
auricular lobules. 34 smooth scales round the middle of the body ; 
dorsals largest, laterals very small. A pair of enlarged praeanals. 
The adpressed limbs just meet. Digits rather short, slightly com- 
pressed ; 18 smooth laraellse under the fourth toe. Dark brown 
above, with strong metallic gloss ; sides with curved or oblique black 
bars ; lower surfaces yellowish. 

millim. 

Total length 166 

Head 19 

Width of head 12 

Body 87 

Fore limb 24 

Hind limb 37 

Tail (reproduced) ()^ 

A single specimen from Faro Island. 

Lygosoma concinnatum, sp. n. 

Habit lacertiform ; the distance between the end of the snout and 
the fore limb is contained once and one fifth to once and two fifths 
in the distance between axilla and groin. Snout short, obtuse; 
supraocular regions swollen. Lower eyelid scaly. Nostril pierced 
in a single nasal ; no supranasal ; a single anterior loreal (except in 

23* 



336 MR. G. A. BOULKNGER ON THE [Mar. 15, 

one of the specimens, which has a very smnll shieLJ ahove it) ; 
rostral foriiiiiig a hroad, strai<;ht suture with the frontonasal, whicl) 
is much broader tlian Ions; ; prpe'roiitiils torminu: a median suture ; 
frontrd much narrowc<l |)0>tcriorl\ , as lonjjas or shorter than fronto- 
pariftiils and iiiter|iarietal together, in cont:ict with the first and 
second supraoculars; four supraoculars, first longest; eight or nine 
supraciliaries, first largest ; frontoparietals and interparietal distinct, 
former lunger than latter ; parietals forming a suture beliind the 
interparietal; no enlarged nuchals ; fiftl) upper Inhial largest and 
beJow tlie centre of the eye. Kar-opening ov;d, nearly as hirge astiie 
eve-opening ; no auricular lobules. Scales smootii, or dorsals and 
laterals indistinctly ))luricarinate, laterals smallest; 40 scales round 
the middle of the bo<ly. A pair of enhtrged pr?e:inals. The hind 
limb readies the elbow of tlie adpressed fore limb or tlie axilla. 
Digits rather elongate, slightly compressed ; subdigital lamellae 
suiooth, 2-' to 25 under the fourth toe. Tail once and a half the 
length of head and bodv. Dark biown ahove, with strong metfiUie 
gloss: back black-spotted; sides witii black and wliitish spots 
elegantly arranued ; a black band on each side of the head, passing 
tiiroi.gli tlie eve ; sometimes a large, blark, vvhitp-ed:;ed spot on eacb 
side of the neck ; lower surlaees brownish white, clouded or longi- 
tudinally streaked with darker. 

millim. 

Total length 130 

Head l^ 

Width of head 9 

Body 49 

Fore limb 17 

Hind limb 27 

Tail (reproduced) 68 

Four specimens from Faro Island. 

Lygosoma albofasciolatum, Gthr. 
Faro Island. 

Typhlops ALTJENsis, sp. 11. (Plate XXVIII. fig. 2.) 

Body much elongate, of subequal diameter throughout. Snout 
depressed, rounded. Nasal comjiletely divided ; a prreocular 
separates the nasal from the ocular, which rests on the third and 
fourth upper labials ; eye very distinct ; the so-called rostral rounded 
posteriorly, its width about three fifths the distance between the eyes ; 
a small azygos shield separates the lostral from the mouth. 22 
scales round the middle of tlie body. Tail comparatively long, twice 
and one third as long as broad at the base, tapering, ending in a 
spine. Brown above, yellowish inferiorly ; upper head-shields edged 
with yellowish. 

Total length 245 millim. ; diameter of body 4 ; length of tail 10. 

A single specimen, from Alu, Shortland Islands. 



1887.] HERPETOLOGY OF THE SOLOMON ISLANDS. 337 

BATRACHIA. 

CORNUFER DORSALIS, A. Dum. 

Faro Island. 

Batrachylodes, g. n. Rauidarum. 

Pupil horizontal. Tongue oval, free and feebly nicked behind. 
No vomerine teeth. Tympanum distinct. Fingers and toes free, 
the tips dilated into large disks. Distal phalanges T-shaped. 
Omosternum and sternum with a bony style. 

Batrachylodes vertebralis, sp. n. (Plate XXVIII. fig. 3.) 

Snout short, obtuse ; loreal region nearly vertical ; nostril nearer 
the tip of the snout than the eye ; interorbital space broader than 
the upper eyelid ; tympanum three fifths the diameter of the eye. 
First finger shorter than second ; disk of third finger as large as the 
tympanum; disk of toes smaller than of fingers; subarticular 
tubercles feeble ; a rather indistinct, oval, inner metatarsal tubercle. 
When the hind limb is pressed against the body, the tibio-tarsal 
articulation marks the posterior border of the eye. Skin smooth 
above and below. Grey-brown above; a fine whitish veitebral line, 
continued along the upper face of the thigh and the outer side of the 
tibia and tarsus; a whitish line on the canthus rostiahs, extending 
from eye to eye; it is continued behind the eye, as a gradually 
widening band, to the groin ; side of head and of anterior half of 
body dark brown ; indistinct brown bands across the limbs ; lower 
parts dirty white. 

From snout to vent 30 millim. 

A sina-le adult female, from Faro Island. 



•»• 



Hyla lutea, sp. n. (Plate XXVIII. fig. 4.) 

Tongue oval, slightly free and very slightly nicked behind. 
Vomerine teeth in two strong transverse groups close together 
between the choanse. Head much depressed, as long as broad or 
slightly broader than long; snout rounded; canthus rottralis very 
indistinct ; loreal region concave ; nostril nearer the tip of the snout 
than the eye, its distance from the latter equal to its dameter; 
interorbital space broader than the upper eyelid ; tympanum very 
distinct, about two thirds the diameter of the eye. Fingers half- 
webbed, the web nearly reaching the disks of the second and third 
fingers; disks larger than the tympanum; no projecting rudiment 
of poUex. Toes three-fouiths webbed, the disks as large as the 
tympanum; subarticular tubercles small and flat; a small, flat, 
inner metatarsal tubercle ; no cutaneous tarsal fold. When the hind 
limb is pressed against the body, the tibio-tarsal aiticulation reaches 
the tip of the snout or a litlle beyond. Skin smooth ; belly and 
lower surface of thighs with large flat granules. Uniform lemon- 
yellow above, white interiorly ; a white line along the outpr side of 
the forearm and fourth finger and of the tarsus and fifth toe. 



338 ON THE MILK-DKNTITION OF THE KOALA. [Mar. 15, 

Male with an internal subgular vocal sac, and black nuptial ex- 
crescences on the inner finger. 

From snout to vent 67 millim. 

Three specimens from Faro Island. 

EXPLAJSfATION OF PLATE XXVIII. 

Fig. I. Lepidodacfylus ivoodfordi, p. 334. 

1 a. . Lower view of foot ; multiplied 3 times. 

2. Typhlops aluensis. p. 33(5. Upper view of head ; multiplied 4 times. 

2 a. . Side view of bead ; multiplied 4 times. 

2 6. . Lower yiew of head; multiplied 4 times. 

2 c. ■ . Lower view of tail. 

3. Bafrachylodes vertebralis, p. 337. 

4. Hyla lutca, p. 337. 



4. On the Milk-dentition of the Koala. 
By Oldfield Thomas. 

[Eeceiyed February 15, 1887.] 

Among the few remaining Marsupials in which no trace of a milk- 
dentition has yet been found, the Koala {Phascolarctos cinereus) 
occupies a prominent place, especially as in this animal the last pre- 
molar, or pm.^, which among Marsupials is the only tooth that 
ever has a milk predecessor, is unusually large and powerful, and 
might have been therefore expected, as in the allied Phalangers, to 
have a proportionally well-developed predecessor. 

At last, however, I have been able to find traces in the Koala of 




Head of young Eoala, showing milk-dentition ; natural size. 



just such a rudimentary milk-dentition as has been described in the 
Thylacine by Prof. Flower \ and showing, just as in that animal, 
that the ancestors of the Koala have had, and that it has now lost, 
the ordinary amount of tooth-change found in the great majority of 
Marsupials. 

In two very young and hairless Koalas, four and five inches long 
respectively, I find, on cutting open the side of the jaw, clear and 

1 Phil. Trans. 1S67, p. 63, 



1887.] MR. G. A. BOULENGER ON A NEW GECKO. 339 

distinct calcified milk-teeth, as shown in the accompanying drawing 
(fig. p. 338). Both above and below they lie in the groove on the outer 
side between the uncut pm/ and m.', their summits being slightly 
above the level of these teeth, but yet not projecting above the gum. 
They are each about 4 millim. in length, the upper one with a conical 
root and thickened crown about 2 millim. in diametei', while the lower 
one is slenderer and has a proportionally longer root and smaller 
crown. 

It is quite evident that these teeth never become functional, but 
are absorbed long before the animal is old enough to be able to 
use them, and in all probability they never cut the gum. 

The discovery of milk-teeth in the Koala is of considerable interest 
when viewed in relation to their comparatively long persistence in 
the Phalangers on the one hand, and their entire absence, so far as 
is yet known, in the Wombats on the other, the Koala presenting in 
this, as in so many other characters, an intermediate condition between 
the two. 

In this connection, however, it may be noted tl>at throughout the 
Mammalia rootless-toothed animals do not have the same need of 
a functional milk-dentition as do rooted-toothed ones, owing to the 
manner in which the ever growing teeth are able to increase in size pari 
passu with the growth of the animal. No better example of this can 
be quoted than the case of the allied Rodent genera Cavia and 
Dasyprocta, the first having rootless premolars, whose milk-teeth 
are absorbed before birth, and the second having rooted premolars 
preceded by well-developed and long-persistent milk-teeth. 

The bearing of this rule on our present subject is evident ; for 
while the entire absence of milk-teeth was quite to be expected in 
the case of the rootless-toothed Wombats, their extreme state of 
reduction in the Koala is a most surprising fact, especially as there 
aie in the latter animal no anterior premolars to make up during 
youth for the absence of milk-teeth, as there are in the Thylacine, 
in which a similar reductiou of the milk-dentition has taken place. 



5. On a new Gecko^ of the Genus Chondrodactylus , from the 
Kalahari Desert. By G. A. Boulengek, F.Z.S. 

[Eeceived March 3, 1887.] 

Mr. J. J. Weir, F.Z.S. , has handed over to me two small Lizards 
from the Kalahari, to be presented to the Natural History Museum 
in case they should prove of interest. Although unfoitunately in 
a dry state, having been pinned in an insect- box, they are in 
comparatively good condition. One belongs to the well-known 
Eremias lugiibris. Smith, the other represents a new Gecko of the 
genus Chondrodactylus, Peters, of which a single species was known, 
C. avgulifer, Peters, also from South Africa. The discovery fa 
second species is therefore of great interest, and I have much pleasure 
in connecting with it the name of Mr. Weir. 



340 REV. G. H. R. FISK ON THE DESTRUCTION OF SNAKES. [Apr. 5, 

Chondrodactylus WEiRi, sp. nov. 

Distinguished from its ally in the following points : — Tubercles 
on tlie supraorbital edge scarcely enlarged, separated from those on 
the other side by three series of tubercles in the middle ; the width 
of the interorbital space equals quite one half of the vertical diameter 
of the orbit. Enlarged dorsal tubercles larger, more strongly keeled, 
subtrihedral. Ventral scales much larger; 6 or 7, on the middle of 
the belly, correspond to the horizontal diameter of the eye (instead 
of 11 or 12 in C. angulifer). Coloration very similar to that of the 
adult C. angulifer, i. e. with a blackish crescentic band, concavity 
forwards, extending from shoulder to shoulder, and pairs of round 
whitish spots on the back. The unique specimen measures 
95 millim., in which the tail enters for 40. 



April 5, 1887. 
Prof. W. H. Flower, LL.D., F.R.S., President, in the Chair. 

The Secretary read the following report on the additions to the 
Society's Menagerie during the month of March 1887: — 

The total number of registered additions to the Society's Mena- 
gerie during the month of March was 7%. Of these 22 were by 
birth, 43 by presentation, 6 by purchase, 1 by exchange, and 4 were 
received on deposit. The total number of departures during the 
same period, by death and removals, was 94. 

The most noticeable additions during the month were : — 

1. Two Long-tailed Grass-Finches {Foephila aeuticauda), from 
Derby, King Sound, N.W. Australia, presented to the Collection by 
Mr. Walter Burton, F.Z.S., March 18. These are the first examples 
of this elegant little Grass- Finch which have been received by the 
Society. 

2. A Fisk's Snake {Lamprophisjiskii) and a Narrow-headed Toad 
{Bufo augusticeps), from South Africa, presented to the Society by 
the Rev. G. H. R. Fisk, and received 24th March. Both of these are 
new to the Society's Collection, and Fisk's Snake, being new to 
science, has been named by Mr. Boulenger after its donor. 

I also wish to call attention to the tact that Sir Walter Buller 
has presented to the Society the female Huia-bird {Heteralocha 
gouldi) which he deposited in the Society's Gardens on the 22nd 
April last year, ami that he hopes to be able to obtain for us a 
companion of the male sex. The female bird in the Gardens is 
now in good health and condition. 

The following extracts were read from a letter addressed to the 
Secretary by the Rev. Geo. H. R. Fisk, C.M.Z.S., dated Capetown, 
March 9, 1S87:— 

"The annexed anecdote of a Mouse^ and a Ringhals Snake (Sepedon 

■' [In a subsequent letter Mr. Fisk states that the Mouse was believed to be a 
specimen of Dciidroiiiyi 77iclanoiis. — P. L. S.] 



1887.] REV. G. H. R. FISK ON THE DESTRUCTION OF SNAKES. 341 

hamachates) was written for me at my request by my friend Mr. 
Sydney Cowper, who, you may remember, was the Cape repre- 
sentative at the late Colonial and Indian Exhibition. His name is 
guarantee of strict accuracy. 

" I send you a copy of his writing, thinking it may be interesting, 
showing as it does a way in which perhaps many young snakes are 
destroyed. Were not an imm.ense number of the eggs and of the 
young of snakes destroyed by their natural enemies, their number 
would soon in some parts become so great as to be very inconve- 
nient indeed to other animals and to man also. 

" I have long known that cats kill snakes. I have seen a lizard 
kill a snake. You will remember a snake which I sent to your 
Society which had devoured the eggs laid by another snake, and now 
we have an instance of a Mouse killing and eating a young venomous 
snake. 

" Probably there are many other ways in which great numbers 
are destroyed before they reach an age and size when they become 
very dangerous. 

" 'On Saturday the 19th February my friend Mr. W. Holms and 
I managed to secure on Wynberg flats, without injury to the 
specimens, two young ' Ringhals,' probably from 7 to 14 days old, 
measuring the one some 10 inches and the other 9 inches in length. 
"We brought them home in our handkerchiefs, placed them in a band- 
box, and proceeded to find food for them. A tour round the garden 
(Rokeby, Wynberg) produced one tortoise, one toad, one field-mouse, 
one cricket, two spiders, and some gentles. These, excepting the 
toad, were all placed in the bandbox with the two snakes, and 
we expected to find the snakes in good condition the following 
morning. 

" ' On looking into the box next morning I found but three sur- 
vivors of the previous night, namely the tortoise, the mouse, and one 
'Ringhals.' The mouse had evidently had the best of it, for he 
was devouring the remains of one of the snakes, and, judging by the 
distention of his little abdomen, I think he must also have con- 
sumed the ciicket, spiders, and gentles. I watched the survivors 
attentively during Sunday, and saw the mouse make an onslaught on 
the remaining Riugiials. He fastened on the snake's back with his 
tiny sharp claws and pecked away with his teeth, the snake trvino- 
its utmost to wriggle away and to secrete itself under the tortoise, 
which it eventually managed to do. The snake seemed much 
frightened, and, although he struck at the mouse frequently, and 
sometimes with apparent success, tiie mouse generally avoided tiie 
stroke with the utmost agility, and before letting go had ridden 
three or four times round the bandbox on the snake's back. I 
imagine that the fang of a young 'Ringhals' is not sufficiently 
developed to penetrate the thick hair on a mouse. I have written 
this account to you, as the fact of the mouse having eaten the snake 
is antagonistic to the generally conceived idea of reptilian customs. 

" 'The Ringhals left for England by the R.M.S. Hawarden Castle 
on the 2nd inst., and the mouse I returned to his habitat under the 
stump of a tree in the garden, and although 1 have several times 



342 MR. F. DAY ON SCOUP^NA SCROFA. [^pr- 5, 

tried to catch hi in again, I have (unfortunately for me) been unsuc- 
cessful. — S. COWPER.' " 

Mr. J. H. Leech, F.Z.S., exhibited specimens of some new 
Butterflies from Japan and Corea, which he was intending to describe 
at a future meeting of the Society, and gave some account of his 
expedition to those countries in quest of Lepidoptera. 



A communication was read from Prof. J. H. Scott and Prof. T. 
J. Parker, containing a description of a Whale of the genus Zip/iius, 
of which a specimen had been recently obtained near Duuedin, New 
Zealand. 

This paper will be printed entire in the Society's ' Transactions.' 



The following papers were read :- 



1, On the Occurrence of Scorpcena scrofa off the South 
Coast of England. By Francis Day, C.I.E., F.Z.S. 

[Eeceived March 26, 1887.] 

On March 21st I was fortunatae enough to secure in Cheltenham 
a recently stuffed specimen of Scoi-pcena scrofa, 11'2 inches in 
length, which had been obtained under the following circumstances. 
It had been captured by a trawler at Brixham at the beginning 
of the month, and forwarded next day to Mr. Woore, fishmonger 
in this town, as being a fish quite new to the local fishermen. 
Owing to my being away and to obviate its being spoiled, Mr. Woore 
had it stuffed, and in this condition I first saw it. So far as I know, 
this fish has not previously been obtained along our shores, and I 
think its occurrence ought to be recorded. 

B. vii. D. 11/yV P. 19. V. i. A. |. C. 13. L. r. 46. 

The specimen agrees so thoroughly with the description in Cuvier 
and Valenciennes's ' Ilistoire Naturelle des Poissons,' vol. iv. p. 288, 
tiiat further remarks upon this subject ap])ear to be unnecessary. 
Although in the ' Catalogue of the Fishes of the British Museum ' this 
species is described as having " the head entirely scaleless and 
smooth," and no mention of spines exists in the description, still in 
the definition of the genus it is remarked that " the head is armed 
with spines." Valenciennes refers to "les nombreuses epines de 
sa tete," and Moreau, in his ' Poissons de la France,' very accurately 
describes the fish. 

Hub. The Mediterranean and along the Atlantic shores of France 
as high as the Gironde and Rochelle. Moreau observed that he had 
never seen it from the coast of La Vendee. Common also at 
Madeira. It does not appear remarkable that a straggler should 
occur along our southern shores, but its occurrence during a very 
cold March would hardly have been anticipated. 



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188/.] ON THE MORPHOLOGY OF THE WINGS OF BIRDS. 343 

2. On some Points in the Morphology of the Wings of Birds. 
By Richard S. Wray, B.Sc. Lond. (Communicated 
by Professor Flower, P.Z.S.) 

[Received March 7, 1887.] 
(Plates XXIX.-XXXII.) 

Since the publication of Sundevall's paper "On the Wings of 
Birds " in 18-43 but little advance seems to have been made in our 
knowledge of the disposition and modification of tlie feathers of the 
bird's wing, although his original Swedish paper was twice translated 
into German. In fact the paper, though forty years old, contains 
much information not to be found in modern descriptions, a great deal 
of it having apparently been overlooked. I have had occasion to go 
into the subject somewhat fully in preparing specimens to illustrate 
the structure of the bird's wing in the Index Museum of the British 
Museum (Natural History). While doing this I found the ordinary 
descriptions unsatisfactory, and at times could not reconcile what I 
saw with them. This occasioned me to examine a great many birds' 
wings of different groups, and led to the results described in the 
following paper. The wings were all examined with a view to make 
out the mode of insertion of the feathers, their relations one to 
another and to the bones, and dried skins were used only when fresh 
specimens were unavailable. Through the kindness of Professor 
Flower I had great facilities afforded me in the way of obtaining 
specimens, and I take this opportunity of expressing my great thanks 
to him for his encouragement and assistance throughout the work. 

While Sundevall's paper gives the correct relations of the parts, 
especially of the coverts, yet many points with regard to the remiges 
and greater coverts he seems to have overlooked, and of others his 
interpretation is erroneous. The relation of the remiges to the bones 
of the manus is not fully described nor accurately figured. That 
the primaries form two groups, metacarpals and digitals, is recog- 
nized, but the absolute constancy of the most proximal digital 
resting upon the phalanx of digit iii. has never been insisted upon ; 
Sundevall's figure shows it as having no connection with the phalanx. 
The presence of one or two more dorsal greater coverts than remiges 
on examination turns out to be erroneous, since every one may be 
accounted for. The presence of a small accessory remex (remicle) 
which I have made out renders the interpretation of the relations of 
the coverts to the remiges more intelligible. These and some other 
important points are discussed in the present paper. References to 
Sundevall's paper are to the English translation which appeared in 
'The Ibis" for October 188G, and are indicated thus (S. p. 39G). 

The nomenclature adopted is founded upon that most in use 

^ " On the Wiugs of Birds," by C. J. Sundevall. Translated from the 
original Swedish of the ' Kougl. Vetensk.-Akad. Handliugar,' 1843, bj W. S. 
Dallas, F.L.S. (Ibis, 1886, p. 389.) 



344 MR. R. S. WRAY ON TtlE MORPHOLOGY [^^PI"- 5, 

at present, and has this advantage that it is appUcahle to both sides 
of the wing, and reduces the terms used to a minimum. Professor 
Flower and Dr. Schiter have done me the kindness of revising the 
nomenclature' for theremiges. The term " tertials " or "tertiaries" 
has been abandoned, " cubitals " always including them when pre- 
sent, because there is no way of absolutely distinguishing any definite 
number of remiges as belonging to this special category. There is 
certainly a distinction to be founded upon the arrangement of the little 
muscular slips and tendons attached to the cubital remiges ; but it 
would not be of much use in practice, owing to the difficulties in the 
way of determining it with regard to many birds. 

The main points of interest brought to light by the examination of 
a considerable number of birds, some of almost every large group, 
will be treated of, the wing of the Wild Duck, which is an extremely 
good type, being first described in detail. The preparations in the 
Natural-History Museum fully illustrate this paper, and most of the 
accompanying drawings are taken from these preparations or from 
essentially similar ones. 

The Wild-Buck's B^ing. 

When the wing is extended for flight, the surfaces and borders 
correspond to those of the primitive vertebrate limb, the preaxial 
border being directed forwards, the postaxial backwards, and the 
dorsal and ventral surfaces upwards and downwards respectively. 
It is in this position the wing is best studied ; and when plucked of 
feathers posteriorly it presents a fold of skin from the elbow to the 
tip in which the flight-feathers and their principal coverts are 
embedded ; these and their position are first described. 

When the wing is prepared as shown in the drawing (Plate XXIX.) 
two main groups of quill-feathers are seen : — the secondaries or 
CUBITALS attaclied to the ulna, and the primaries or metacarpo- 
DiGiTALS attached to the manus. Of the latter, six, the Meta- 
carpals (l-ti), are attached to the metacarpus, and five, the Digitals 
(7-11), attached one {acldigital, 7) to phalanx 1 of digit in., two 
(middigitals, 8, 9) to phalanx 1 of digit ii., and two {predigitals, 
10, 11) to phalanx 2 of digit ii. The distal predigital (11) is 
always small, and is designated the remicle ; its relations, described 
in detail later, show that it is as much a primary as the so-called 
" spurious tenth " of many Passerines. The quill-feathers on tlie 
cubitus stand out more or less at a right angle to the bone ; those on 
the manus form a gradually increasing obtuse angle, till the last 
feather lies parallel with the phalanx to which it is attached. 

The remiges are best numbered from the wrist-joint, proximally 
for the cubitals and distally for the metacarpo-digitals ; because 
with scarcely any exception reduction in number takes place at the 
distal end of the manus and the proximal end of the cubitus. 

^ A somewhat similar nomenclature was proposed by Dr. Alix, ' Jouraal cle la 
Societe philomatique,' 1874, p. 10. " Sur les plumes ou remiges des ailes des 
oiseaux." 



18S7.] OF THE WINGS OF BUIDS. 345 

The remainino; feathers of the wing are the coverts; thev are best 
understoml if described frnin the posterior mariiin of the vvina;. 

Oil tlie dorsal side the row of feathers (Plate XXX. fij:. a, n) 
Ivinj;; next the remises are the tuctrices majorcs, being quite definitely 
related to the reinigeal quills, and lying close pressed upon their 
bases. Each remex is serial with the covert proximal to it, the 
cubital coverts crossing over tlie bases of the rfniiges, the meta- 
car[)al coverts lying parallel and pressed upon two contiguous re- 
miges (c/l Plate XXXI. figs, b and c). There is a well-developed 
covert to every metacarpal exeeut tiie first, which possesses only a 
very small and vestiu;ial one, 1', vvhieh is completely hidden by a 
median covert, 1" (Plate XXXII. fig. 8), which in many birds func- 
tionally replaces it, the t. ma/or disap[)e.iring. 

On the ventral surface of the wing is a row of featliers (Plate XXX. 
fig. b, «), bearing the same relations to the reniiges as those just 
described ; these are the (ectrices majores of tlie lower surface (e/'. 
Plate XXX. fig. 6, and Plate XXXI. fig. b). If fig. b, Plate XXX 11., 
be examined, which shows the relations of the above feathers in 
section, starting at thf tip of the wing tiie remicle, or jiredigiial 2 
(R'), is seen to have proximal to it a dorsal and ventral covert, term- 
ing a group of three. The next remex is similar, and so to the 
1st metacarpal, whose dorsal covert is very small and rudimentary. 
These relations show that the remicle is a small metacarpo-digital 
whicli has probably not been differentiated into a flight-feather. The 
cubitals show the same arrangement except the fiftli group, where 
there are a pair of coverts, but no remex ; this condition is termed 
aquincubital, and is later described more fully. 

On the dorsal surface the next row of feathers to the t. majores 
are the median coverts (Plate XXX. fig. a, /3), or tectrices medicp, 
arranged serially with the other groups. On the cubitus they lie 
with a reversed overlap to the remiges and t. majores ; those, how- 
ever, which lie most proximal are unreversed (S. p. 4\f>, footnote, 
and Goodchild, P. Z. S. 1880, p. 191). Those on the manus lie 
unreversed, and generally the median covert of the 2nd metacarpal 
is wanting (c/". Plate XXXII. fij:. 8). On the ventral surface of the 
wing, the next row of feathers (Plate XXX. fig. b, /3) bears similar 
relations ; they are the tectrices media of the lower surface, and 
always lie with reversed overlap to the remiges and t. majores (S. 
p. 491). The distal tour or five are generally deficient on the manus 
in the Duck (Plate XXXI. fig. b). In many birds they are nearly 
all suppressed on the manus. 

The tectrices majores and mediEc on the ventral surface have at 
first sight an anomalous position. Being on the ventral side of the 
adult wing, one would expect the backs of the feathers to look 
ventralwards, whereas they look dorsalwards just as do the remiges. 
This is pointed out by Sundevall (S. p. 419), who, however, gives au 
erroneous explanation, saving they are aftershafts developed at the 
ex >ense of the true feather-shaft; a more probable explanation is 
discussed later. 

The feathers so far described are seated in the wing-membrane, 



346 



MR. R. S. WRAY ON THE MORPHOLOGY 



[Apr. 5, 



the next rows being in the skin covering the muscular portion of the 
wing and in the patagium. On the dorsal surface five rows of 
feathers (Plate XXX. fig. a, y) follow the t. medise, lying with the 
same overlap, and on the manus being scantily represented ; they 



Fig. 1. 



© 6 




L © ^' 7 




12 3 



n.c 




VI 

a, a'. Drawings of preparations of the distal cubital remiges, Tvitb tbeir attached 

tectrices majores, of the Pheasant. 

a. Dorsal view; a', ventral view. (This shows the " quincubital" condition.) 

h, h'. Drawings of preparations of the distal cubital remiges, with their attached 
tectrices majores, of the Golden Eagle. " Aquincubital." 

b, Dorsal view, b', ventral view; 1, 2, 3, &c. the remiges (/?), numbered from 

the wrist-joint ; B.C, dorsal tectrix major; V.C, ventral tectrix major; 
Til, ulna. 



I 



are the tectrices minores. They extend on to the arm, and on tlie 
dorsal surface of the humerus a row of 6 feathers becomes elongated, 
forming an apparent continuation of the remiges of the forearm, 
the feathers of the next row taking the form of coverts ; they 



1887.] OF THE WINGS OF BIRDS. 347 

form the humevals (pennce humerales), the "parapteron" of Nitzsch 
(Plate XXX. fig. a, h). On the ventral surface next the median 
coverts are three row of feathers (Plate XXX. fig. b, y), the tec- 
trices minores of the lower surface, which are but scantily repre- 
sented on the manus. 

A fairly well-marked space ' running the whole length of the 
cubitus separates these from two to three rows of feathers which run 
from the wrist to the elljow ; and then are continued on the arm, 
where they become largely developed, 6 to 8 feathers (Plate XXXI. 
fig. b, ax) forming the axillaries (hypopteron of Nitzsch). This row 
may be termed, when distinct as here, an axillary row, though really 
forming part of the minores ; in some birds there is no space sepa- 
rating them, and then they are confluent. The t. minores of the 
upper and lower surface generally correspond, both producing special 
developments, the " humerals " and " axillars." 

The next group of feathers (Plates XXX. & XXXI., /i) grow along 
the posterior border of the wing, extending from the proximal end of 
the patagium to the end of the manus. On the dorsal surface they 
soon approach the minores and become confluent with them, though 
distinguishable in fresh undisturbed plumages by difference of over- 
lap. On the ventral surface the patagial space is large, and separates 
them well from the minores. The feathers growing from the edge 
of the patagium are sufficiently elongated to cover this deficiency. 
At the wrist they become confluent with the other series and are 
• continued on the hand. This group of feathers is common to both 
surfaces of the wing, insomuch as they form on the anterior border 
a shelving series, giving a clean finishing edge to the anterior 
margin of the wing. Tbey are best termed marginals (fecirices 
marginales). The feathers of the pollex, pluma pollicis (" alula," 
"ala spuria"), are partly of this series and of the minores ; and by 
specialization produce four small quill-feathers with coverts, which 
lie closely embracing the dorsal part of tlie anterior border of the 
manus, and hiding many of its lesser coverts. 

The table at the end of this paper (p. 355) shows the relation 
of the nomenclature adopted above with that of Sundevall, and the 
ordinary nomenclature such as that found in Coues's Key to N. A. 
Birds. All birds' wings (except the Penguins) are directiv referable 
to the type just described. It contains all the elements which occur 
in the wing, and it is by the specialization and suppression of these 
parts that the different wing-forms have been derived, at any rate 
among Carinates, the Ratite wing being more primitive in structure. 

Some Modifications of the Wing. 
The remiges of the manus show a remarkable constancy both in 
number and fiosition, for (with the sole exception of the Penguins) 
tiie first digital always lies upon the phalanx of digit iii., its end 
resting upon the metacarpo-phalangeal articulation ; the middigitals 
are always constant in position, so nve the predif/itals. On the meta- 
carpus are six feathers always except in Flamingoes, Grebes, and 
^ This is especially well seen in the Grebes. 



348 MR. R. S. WRAY ON THE MORPHOLOGY [Api*- 5> 

Storks, where seven occur. The prerlic/itrils are the only other 
reini<;es of the manns wliicli show modifications of any interest. In 
tlic typical condition (cf. Plate XXXI. fia;. «) we have the lar;;e pre- 
dijjital 2 (a) and the small remicle (/3), witli their dorsal and ventral 
coverts all intimately attached to the phalanx. Tiiis arrangement is 
probably generally present in the Pygojiodes, Gaviae, Tubinares, 
man}' Limicolse, Flerocles, Odontoglossse, Herodiones, Anseres, 
Pelicans, Siriges, and Accipitres. Among other birds it is probably 
not genernllv present, but it may be found in many of tiie lower 
forms of Passeres, and in some Picarise, prcibably never in Galiinae. 

The remicle disappears in these forms, but its dorsal covert 
remains well develojied, especially well seen in the Galiinae, and its 
ventral covert may also remain, but often disappears. All trace of 
tiie •iroup may disappear, as in tlie nine-primaried Pas-erines, where 
predigital 2 is reduced to a mere rudiment, but can generally be 
detected ; its covert is always well marked. The so-called pencil- 
feather of tlie Woodcock is the dorsal covert of the remicle group. 

Tb.e chief, most interesting, and most puzzling modification of the 
cubital feathers is that in a great many birds the fifth remex is 
always undeveloped, its coverts being normally developed and present. 
This occurs probably in all birds except Phcenicopterus, Galiinae, 
Passeres, and a few Picarise. Up to the present I have never met 
with a trace of this feather in a vestigial condition. If the Hsures of 
the preparation of the distal part of the cubitus of the Golden Eagle 
be compared with those of the Pheasant (see p. 346), the exact nature 
of this modification is at once apjiarent. In the Pheasant («, a') the 
fifth remex is present with its coverts, showing all normal relations; 
in the Golden Eagle {b, b') the coverts are present but no remex. 
The former condition may be termed quincubital, the latter aquin- 
cubital. Such is the constancy of one or tlie other condition in each 
natural group, that I have as yet met with no exceptions anywhere, 
except among the so-called Picarice, many of which are, and most 
of which probably will (urn out to be, quincuhital. The Goatsuckers 
are aquincubital, while the Swifts are quincubital. Pterocles is 
aquincubital ; Goiira is aquincubital. Of course exceptions may turn 
up, seeing that of the whole number of birds but a comparatively 
few have as yet been tested for this point. 

In the Galiinae the first cubital feather is shortened ; this is 
possibly due to mechanical requirements in the folding of the wing, 
as the metacarpal remiges are inserted so near the actual joint as to 
leave but little room. Nitzsch states that sometimes the last feather 
on the mauus undergoes shortening. I have not met with this 
condition. 

In the description of the Duck's wing it was pointed out that the 
upper major covert to tlie first metacarpal remex is very small and 
rudimentary. When the feathers are all plucked off except the 
remiges, major and median coverts, the appearance at the wrist- 
joint is that represented in the figs. 6-9 (Plate XXXII.), where the 
remiges are red, the major coverts yellow, and the median blue. The 
diagram above each of the figures shows the real homologies of these 



1887.] OF THE WINGS OF BIRDS. 349 

feathers, in their undisturbed primitive conditions. Sundevall men- 
tions that there are generally one or two extra major coverts connected 
with the cubital series, of which the feather / in his figures (the 
median covert here under consideration) is one. Tlie other he does 
uot mention (S. p. 414, par. 2). Really there are no extra coverts 
at all, unless the fifth cubital coverts in the aquincubital condition 
of the wing are so considered. In the Duck (Plate XXXII. fig. 8) 
it is seen that the larger feather (1"), which at first sight appears to 
represent the major covert, is really the median covert in front of the 
remex in the undisturbed quincunx series; the little feather ( 1') under- 
neath being the real major covert. In this wing the two feathers 
have not become so closely attached to the first metacarpal as is 
the case in some other birds, example the Golden Eagle, Barn-Owl, 
&c. (Plate XXXII. fig. 6). Here the relations are more apparent; 
the major covert is small and rudimentary, but bears the same 
relation the other major coverts do. The median covert is a fairly 
large feather, which crosses the metacarpals at a considerable angle. 
In the Duck the median covert of metacarpal 2 is suppressed, in the 
Eagle it is present (2", fig. G). The Grebe, with seven metacarpals, 
shows similar modifications at the wrist-juint (Plate XXXII. fig. 7). 
In many birds {e.g. Passerines) the major covert altogether disappears, 
the median covert is large, and takes its place, lying across the 
metacarpals at a considerable angle, and causing the suppression of 
the next one or two mediae (Plate XXXII. fig. 9). 

The remaining feather-tracts undergo modifications in different 
groups, which are generally of more or less minor importance. The 
axillars and humerals vary in their development in different groups, in 
the Passerines disappearing almost absolutely, in all probability in 
connection with the relative shortness of the humerus. The marginals 
in many birds of this group are much elongated on the ventral side 
and cover the patagium. 

When the wing is folded these feathers (often with the last two or 
three major cubital coverts) present much the appearance of the 
axillars in some birds. They have often been erroneously so described, 
whereas they have nothing whatever to do with them, the true 
axillars being represented by but a few semiplumes at most. 

The modifications of the overlap of the dorsal cubital median and 
minor coverts have recently been very fuUv worked out bv Goodchild 
(P. Z. S. 1886, pp. 184-203). These feathers are'termed by 
Goodchild the " median cubital coverts ;" but it is much better to 
confine the term " median coverts " to the row following the majores, 
and call the others " minor coverts." Goodchild's terms " sup- 
plementary row of median coverts" or " upper wiug-coverts " and 
" posterior row of median coverts " are unnatural, because part of 
his supplementary row in the majority of cases belongs to tbe 
t. mediae, and the greater and proximal part of his posterior row to 
the same, whereas the distal part belongs to the first row of minores 
{cf. his fig. 1, p. 186, P. Z. S. loc. cit.). His supplementary row 
generally means the distal t. mediae proper, and sometimes includes 
some of the feathers of the next two or three rows on the wrist, 

Proc. Zool. Soc— 1887, No. XXIV. 24 



350 MR. R. S. WRAY ON THE MORPHOLOGY [Apr. 5, 

which appear, as a consequence of folding superficially, to form a series 
with them ; it is sim])ly accommodation, and is only very striking 
when the wing is examined in the folded condition, as most of his 
were. Grouping the feathers according to tlieir insertion, and 
remembering the conditions of folding which mnst occur at the wrist, 
the observations of Groodchild give the most complete view of this 
subject we have. The Passerine birds possess only the single row, 
t. media, the minores being completely absent (S. p. 415, cf.); 
this is characteristic of them, and goes along with a very scantily 
feathered ventral surface. 

The median and minor coverts of the lower surface * show a great 
variety of modification, which if systematically worked out would I 
believe furnish valuable characters. But jiractically ornithologists 
take no account of the lower surface of the wing, and but little can 
be made out from ordinary skins. For skins to be of much value 
for studying wing-characters, some of them ought to be prepared 
with one wing in the extended position. Fresh wings are much the 
most valuable, but they are not always available. According to 
Sundevall the median coverts often show a tendency to disappear, 
which I have noticed, often they are very small. In the Passeres the 
row of t. majores disappears ; this is recognizable by difference of 
overlap. 

The Dueldinff's Wing. 

If the wing of a Duckling be examined when it is a mere downy 
appendage, of no use for flight, it will be found to be an exact fore- 
shadowing of the flying wing. The fifth cubital remex is absent, not 
even a vestige of it can be found ; so we may conclude that this 
modification is a very ancient and deep-seated one. The plumules 
(down) clothing the wing are more feather-like than the adult down. 
In fact a Duckling's wing forms an interesting comparison with an 
Ostrich's or Rhea's, these wings probably never having got beyond 
a stage ])arallel to this. The way the plumules of the Duckling are 
shed is very interesting; at the base of theplumule the new pennaceous 
feather forms, grows, and begins to force its way out from the skin. 
The plumule remaining attached by its base to the tip of the new 
feather is carried out away from the skin about i to | inch, then the 
connection becomes very slight, and soon the plumule is lost. This 
forms an analogous parallel to the shedding of milk-teeth, the plumule 
being retained till the new feather can functionally take its place. 

The Wings of the Ratitae. 

The wings of the Ratitae conform to the same general plan as those 
of the Carinatse, presenting a modification of a more generalized type, 
which correlates with their bony structure. 

In the unplucked Ostrich wing, little beyond a confused mass of 
feathers can be made out. The ventral surface is totally devoid of 

^ There are some yery good figures of the lower surface of the wing of 
certain Hawks in the' Zoologist,' 1880, p. 273, pis. 2 and 3. 



1887.] 



OK THE WINGS OF BIUDS. 



351 



Vis. 2. 



Mr I 



Mc2 
Phi 1 Ph.2 

jClau) 




a. Preparation of the manus of the Ostrich, showing the primaries and the 

manner of tlieir attachment to the bones. 

b. Dorsal view of the antebrachium and manus of the wing of the Ostrich.' 
e. Ventral view of the manus of an euibrjo Ostrich. 

M, metacarpals ; B, digitals ; Ad, addigitals ; Md, middigitals ; Pd, predigitals ; 
Cu, cuneiforme; Mc 1, 2, 3, meiacarpals 1, 2, 3; Ph 1, 2, 3, phalanges 
1, 2, 3; I, II, HI, digits i, ii, iii ; T.Mj, tectrices niajores; T.Md, tec- 
trices mediae ; T.JSIn, tectrices minores ; M, marginals ; Al, penna; pol- 
licis (alula) ; 1, beneath this fold are the two ventral t. majores mentioned 
in the text; a', tectrices majores inferior. 



feathers except for one row, the t. majores of the lower surface. If 
the feathers be cut short, so that only the base of the quill is left 
in the skin, their arrangement can then be satisfactorily studied 
{cf. fig. 2). On looking at the figure (2, a), the reniiges, It, are seen 
in their natural position, next a row of t. majores, T.Mj, then the 

24* 



352 MR. R. S. WRAY ON THE MORPHOLOGY [Apr. 5, 

t. medise, T.Md, but scantily represented on the manus. There is 
])art of a row, T.Mti, representing the niinores, and a few roAvs of 
marginals, M. The dorsal surface of the lurmerus is uniformly 
covered by rows of feathers. The pollex bears four rerniges and a 
few coverts, Al. 

The disposition with regard to the bones gives 16 primaries or 
metacarpo-digitals, and about 20 (20-22 or 23) cubitals. The quills 
have not the same firm attachment .<»s those of the Carinates, there 
being no grooves in the phalanges to receive them, and their bases 
project beyond the anterior edge of the bone (e/. fig. 2, p. 351). In the 
Carinatse the quills attached to the phalanges lie almost parallel to 
them, whereas here the angle is little larger than a right angle. This 
is a much more primitive condition. 

The primaries are disposed as follows : — Eight metacarpals, one 
addigital, four niiddigitals, and three predigitals. This probably 
represents a more primitive wing-form than the Carinate, where 
seven metacarpals and five digitals is the highest number of pri- 
maries. Probably tlie ancestral wing-form became modified into 
the forms we know by reduction and specialization of these feathers, 
seen more numerovis in the Ostrich than elsewhere. 

The Rhea's wing presents the same general characters as the 
Ostrich ; the ventral surface is bare, and the dorsal surface, with tlie 
feathers cut, shows the same arrangement ; but when the relations 
of the remiges to the bones are considered, it is seen to approach 
more nearly to the Carinate type in some respects. The primaries 
are twelve in number, there being seven metacarjjals, one addigital, 
two middigitals, and two predigitals. This reduction is correlated 
with shortening and reduction of the mauus. The angle of insertion 
of the digitals is more obtuse than in the Ostrich. 

The wing of the Emu I have not had the opportunity of dis- 
secting, but it is probably similar in arrangement to the Ostrich and 
llhea, judging from a stuffed specimen. 

The wing of the Cassoway ' shows a great exaggeration of the 
feature, noticed in the Ostrich, of the quills projecting beyond the 
bones, its quill-spines being the sole remains of the cubital remiges. 

The Apteryx shows, as was first pointed out by Prof. Flower 
(Roy. Instit. Lect. 188G), a few true cubital remiges, indicated by 
their long quills. 

The PenguirCs Wing. 

This departs the most of all wings from the general plan. The 
paddle form of the wing and its scale-like feathers are familiar, and 
there is little or no differentiation apparent beyond the passage from 
mere scales anteriorly to feathers posteriorly. On the ventral side 

^ In the wing of a Cassowary dissected since writing the above there are 
to be seen structures representing, in ;ill probability, the " primaries," wliich 
appear at first sight to be entirely wanting in these i'orins. I hope to describe 
this specimen, together witli some other interesting Ratite flings, in a future 
paper. 



1887.] OF THE •WINGS OP BIRDS. 353 

this is all. On the dorsal the first four rows of feathers show a 
certain amount of differentiation, being somewhat elongated, and 
showing what might be looked upon as a tendency to form remiges 
and coverts, which was early lost, the wing taking a different func- 
tion to those which developed into organs of flight. The embryo 
of the Penguin shows in its wings no signs of being a degeneration 
or modification of the specialized flight-wing of other Carinates. 
There appears to be no trace of remigial structure at all in this 
wing. 

Origin of JVing and General Conclusions. 

The study of the wings of living birds leads to the conclusion that 
the power of flight was gradually acquired, and also tends to throw 
some light upon the way wings were originally evolved from a 
reptilian manus. Recent researches ^ seem to show that the ances- 
tral form of the avian manus was probably a webbed form, and 
inferentially belonged to an aquatic type of animal. From this 
'^ webbed paw " was developed the starting-point of the wing, by 
special modification of the scales or feather fbretypes on the dorsal 
surface. The Penguin's paddle represents, perhaps, a highly modi- 
fied survival of this starting-point ; the Ratite wings are modified 
conditions of the intermediate stage in the wing-formation. At 
some future time I hope to bring forward the evidence in favour (or 
otherwise) of this view more fully worked out; however, the follow- 
ing are some of the points which tend to support that view. 

In the adult flight-wing of the Carinates there are two rows of 
feathers situate on the ventral side of the wing, reversed in position, 
the t. majores and mediae. Sundevall explains this by saying it is 
an aftershaft developed at the expense of the feather-shaft, and 
states (S. p. 419) that the aftershaft is entirely deficient ; but in a 
Pheasant I have found it normally developed, though small in these 
feathers. His explanation is erroneous. The true explanation 
probably is that these feathers or their antetypes \yere originally on the 
dorsal surface and have been carried down to the ventral in the for- 
mation of the " ala membrana" by the excessive development of the 
remiges and tectrices majores. That is, that originally on the dorsal 
surface of the arm and manus there took place a special modification 
of the scales or feather foretypes by which rows of these were 
directed backwards in the "primitive embryonic" position of the 
limb. Next two or three rows began to be specialized and to become 
larger and more prominent than the others ; then these, by their 
unequal growth, carried over a fold of skin and formed the wing- 
membrane, carrying some of the structures to the ventral side, which 
are now seen as the reversed feathers {cf. diagrams, Plate XXXII. 
figs. 1-5). In the embryo bird the feather-rudiments first appear on 
the dorsal surface, pointing to the fact that the modification here is 
very ancient and deep-seated ; the remiges and greater coverts 
(superior) being the earliest to appear; quickly they begin to assume 

^ Prof. W. K. Parker's recent paper " On the Morphology of Birds," read at 
the Royal Society, Jau. 27, 1887. 



351 MR. R. S. WRAY ON THE MORPHOLOGY [Apr. 5, 

larger proportions, and at the very earliest stages the remiges are dis- 
tinguishable. At this stage the wing is quite rounded in section, there 
being no trace of the " ala membrana ;" the next feathers to appear 
are the t. majores (inferior), closely followed by the other ventral 
coverts, the other dorsal coverts meanwhile having appeared. At 
this stage {cf. Plate XXXII. fig. 1) the inferior major and median 
coverts are distinctly more on the dorsal half of the rounded edge 
of the wing than its ventral, but very quickly they become quite 
ventral, owing to the rapid growth of the remiges. This stage 
is quickly passed over, but sufficient is visible to sliow that these 
feathers are carried distinctly to the lower surface by inequality 
of growth {cf. Plate XXXtt. figs. 2-4). The feathers resulting 
from these are the plumules seen in the wing of the DiickHng, and of 
no use for flight. The wing of a Duckling reproduces in a great 
measure, allowing for specialized differences, the adult Ostrich's 
wing or the Rhea's ; and these wings are survivals of the transition 
state of the wing, probably never having been used for flight, but 
having undergone special modifications of their own from that point. 
It is pretty clear the remiges of the Ostrich and Duck's wing corre- 
spond, more so the Ostrich and Duckling's ; in the Ostrich we have 
but one row of ventral coverts, and in the embryo we get them most 
distinctly on the dorsal side. The Ostrich embryo figured (fig. 2 c, 
p. 3.51) shows the manus from the ventral surface ; digits i., ir., and 
III. being well developed ; digit iii. at its tip projecting beyond the 
general fold of the wing ; in fact there is a very complete webbed 
manus. The feathers seen (a', fig. 2 c, p. 35 1) are the row of ventral 
coverts, and lying over digit in. on its dorsal surface are two of this 
row hidden from sight by it. In the adult, one of these feathers grows 
over the distal part of phalanx 1 of digit in., owing to elongation of 
its quills ; here we have the dorsal position actually preserved in the 
adult. The wing of the Ostrich presents also a primitive condition 
especially in the cubital region, in that the " ala membrana" is not 
specialized as in the Carinatae, being in the intermediate condition of 
the Carinate embryo. Probably the feathers now representing the 
remiges and the principal coverts were more numerous in the primi- 
tive wing type, and iiave become restricted in number on the manus ; 
thus the Ostrich has 16, the Grebes 12, while most birds have only 
1 1 primaries. 

Wing-Formula. 

The main facts with regard to the feathers of a bird's wing may 
be expressed as a formula. Denoting the metacarpo-digitals by 
Md, the metacarpals by m, the digitals by d, and expressing the 
number of feathers in each group by a number placed after (thus, 
six metacarpals, m 6), the cubitals by C, " quincubital," "aquin- 
cubital " by C% we formulate the remiges 

MdU m6 db C'x*. 

The coverts are indicated by o, /3, y, for the t. majores, mediae, 
and minores respectively ; by placing a figure below the line, thus n^, 

* 0' = number of cubitals, wbicb varies considerably in different groups. 



188/-.] 



OF THE WINGS OF BIRDS. 



355 



Ventral Coverts. 
Tectrices inferiores. 



Remiges. 



Dorsal Coverts. 
Tectrices superiores. 



2 

B 



o 



B' 






H 


H 


GO 


hij 


05 


O) 




T 


o 


o 












3 






C3 
CI- 






ffl 




Q 


P 






CO 


1-^ 












a 


3 


2 


o 


a> 


P 




Cl4 

S' 


O 

0) 
00 


o 

►5 


g 




Q 


fp 






o- 


(15 






03 


■^3 
















1 



CD 






2 

o 

1=: 



p 
S' 



a- 2. 



23 

B 
8 



c 



w 


w 


Q 


o 


3 


B 






15 


Uq 


re 


cc 




O) 


o 


*ri 


c 






3 


f=, 


p 



H 


H 


Q 


CO 


O 








"-J 


rj 






ft 


n 


(T> 




09 


y— s X 




CD 3 




S G" 


fe 


g§ 






hi 


cr to 


K 


E o 




ii 


ro' 




p ^. 








0) ■< 




QQ (ti 












B 








t3 




C 




CO 



> 
ca 
f 

H 
O 

o 

H 
2! 
O 

33 



o 

b 

C 



Is- 

a 'I 
op S 

o 




i^s 



13 



Q 

a 

s 
»■ 



356 MR. R. S. WRAY ON THE MORPHOLOGY [Apr. .5, 

it indicates the number of rows, and placing these symbols above or 

below a line, thus ^:jJ^ their dorsal or ventral position ; the mar- 

ginals are expressed by ju, the humerals by h, the axillars by .r, a 
number after the h or'.r denoting the number of specialized feathers 

forming the "parapteron" and " hypopteron." When any two 

— '—, 
rows are confluent it may be indicated thus, yx, where tbe axillars 
and minores are indistinguishable as separate groups ; Al expresses 
the quill-feathers of the pollex. 
Formula for the Duck : — 

Mdll mQdb "jlllA (Pl9^^^y,A±IL AM. 
a,/Ji7/i "i Pi 73 ^ 7(2-3) /* 

Typical Passerine formula : — 

MdlO mQ di "jAyiL C9 "'^'>'°^ A13. 
"i 1^0 7A* «o Pi 7-2 A* 

Formula for Ostrich wing : — 

Md 16 7)1 8 d8 ^i& C20^^^jA1i^ AM. 
«i "i 

These formulae might prove of value to ornithologists by enabling 
them to briefly express the main characters of the wings of diiferent 
groups of birds. The three given above at once express very great 
differences in the wings of these birds : thus it is seen at once how, 
in the Passerine, the upper minores, the axillars, and humerals are 
absent ; and the whole of the lower coverts, except one row, in the 
Ostrich. These are here introduced to show the possibility of using 
a wing-formula expressing most of the characters. 



DESCKIPTION OF THE PLATES. 
Plate XXIX. 

Drawing of a preparation of the right wing of the Wild Duct, seen from below, 
sbow ing the relation of the quill-feathers to the bones. 

al. PlumaB pollicis (alula). C. Cubitals or secondary remiges. Md. Mel,a- 
carpo-digitals or primary remiges. 3L Metacarpals. D. Digitals. Ad. 
Addigital. Md 1 & 2. Middigitals 1 and 2. Pd 1. Predigital 1. Pd 2 or 7?. 
Predigital 2 or remiole. H. Humerus. E. Kadius. Ul. Ulna. 8c. Scaphoid. 
Cu. Cuneiform. Mc 1, 2, .3. Metacarpals 1, 2, 3. Ph 1, 2, 3. Phalanges 1, 
2, 3. I., II., III. Digits I, 2, and 3, 

Plate XXX. 

a. Plan of the arrangement of the feathers on the dorsal surface of the extended 

left wing of Anas boschas. 
h. Plan of the arrangement of the feathers on the ventral surface of the ex- 
tended right wing of Avas hoschas. 
C. Cubitals (grey). Md. Met^acarpo-digitals (grey), a. Tectrices majores 
(pink). /3. Tectrices media; (green), y. Tectrices minores (brown). /<. Tec- 
trices marginales (yellow). Al. Pluma3 pollicis (red). h. Humerals. x. 
Axillars. x'. Axillary row of minores. 



1887.] OF THE WINGS OF BIRDS. 357 

Plate XXXI. 

a. The distal phalanx of digit ii. of the wing of the Barn-OwI, with the attached 
predigitals and their coverts, showing the remicle and its relations. 

1. Ventral view. 2. Dorsal view. 

a. Predigital 1. |8. Predigital 2 (remicle). 

a'. Dorsal tectrix major to a. 

a". Ventral tectrix major to a. 

(3'. Dorsal tectrix major to remicle. 

/3". Ventral tectrix major to remicle. 

1. Phalanx 2 of digit ii. 

2. Fused phalanx 3 of digit ii. 

b and c. Diagrams of the ventral and dorsal surfaces of the wing of the Wild 
Duck, showing the points of insertion of the feathers of the different 
groups. 

C. Cubitals. M. Metacarpo-digitals. i?. Eemicle. r. Remex. 

a. Tectrices majores. 

fi. Tectrices mediae. 

y. Tectrices minores. 

fi. Tectrices marginales. 

.r. Axillars. 

h. numerals. 
Ax. Axillary row. 
(5). Absent fifth cubital remex. 
Al. Alula. 

Plate XXXII. 

Figs. 1-6. Diagrams showing how the ventral tectrices raajores and mediae have 
been carried over from the dorsal side, and the " ala membrana " 
formed. 

1. The earliest condition of the feather-rudiments. 

2-A. Intermediate conditions. 

5. The condition in the adult wing. 

Eemiges — red. 

Tectrices majores (superior) — yellow. 
Tectrices majores (inferior) — green. 
Tectrices media; (superior) — dark blue. 
Tectrices media; (inferior) — light blue. 
These diagrams represent sections across the wing in the direction x-?/ (fig. 8). 

Figs. 6-9. The remiges and upper principal coverts in the region of the Avrist- 
jointin Barn-Owl ((5), Grebe (7), Duck (8), and Lark (9). 
1, 2, 3. 4. Metacarpal remiges (red). 
1', 2', 3', 4'. The corresponding major covert (yellow). 
1", 2", 3", 4". The corresponding median covert (blue). 
* Wrist-joint. 

The plan of these feathers in section is shown above each, and represents the 
primitive unmodified relations. 

a. Drawing of section through the large ^feathers of the wing just below the 

edge of the " ala membrana" of the Pheasant. 
P, The same of tJie Duck. The proximal cubitals are not shown. 
B. Remex (red). 
DC. Dorsal covert, tectrix major (yellow). 
VC. Ventral covert, tectrix major (green). 
i?'. Remicle. 
* The wrist-joint. 
Q. Fifth cubital remex present. 
Aq. Fifth cubital remex absent. 



3.")8 REV. H. S. GORHAM ON THE CLASSIFICATION [Apr. 5, 

3. On the Classification of the Coleoptera of the Subfamily 
Languriides. By the Rev. H. S. Gorham, F.Z.S. &c. 

[EeceiTed March 26, 1887.] 
Family Erotylid^. 
Subfamily Languriides.